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The CANADIAN 
FIELD-NATURALIST 


Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada 


Volume 118, Number 1 January-March 2004 


125" Anniversary of the Ottawa Field-Naturalists Club 1879-2004 


The Ottawa Field-Naturalists’ Club 


FOUNDED IN 1879 


Patrons 
Her Excellency The Right Honourable Adrienne Clarkson, C.C., C.M.M., C.D. 
Governor General of Canada 
His Excellency John Ralston Saul, C.C. 


The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to 
encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields 
as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environments 
of high quality for living things. 


Honorary Members 


Edward L. Bousfield Bruce Di Labio John A. Livingston E. Franklin Pope 
Donald M. Britton R. Yorke Edwards Stewart D. MacDonald William O. Pruitt,, Jr. 
Irwin M. Brodo Anthony J. Erskine Hue N. MacKenzie Joyce and Allan Reddoch 
William J. Cody John M. Gillett Theodore Mosquin Mary E. Stuart 
Francis R. Cook C. Stuart Houston Eugene G. Munroe John B. Theberge 
Ellaine Dickson George F. Ledingham Robert W. Nero Sheila Thomson 
2004 Council 

President: Mike Murphy Ronald E. Bedford Barbara Gaertner Stanley Rosenbaum 
Vice-President: Gillian Marston Fenja Brodo Diane Lepage Louise Schwartz 
Recording Secretary: Susan Laurie-Bourque John Cameron Christina Lewis David Smythe 
Treasurer: Frank Pope William J. Cody Karen McLachalan Hamilton Henry Steger 
Past President: Gary McNulty Kathy Conlan David Hobden Chris Traynor 

Francis R. Cook Cendrine Huemer 


To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate 
P.O. Ottawa, Canada K1Z 1A2, or e-mail: ofnc @achilles.net. 
For information on Club activities telephone (613) 722-3050 or check http//www.achilles.net/ofnc/index.htm 


The Canadian Field-Naturalist 


The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this 
journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. 


We acknowledge the financial support of the Government of Canada toward our mailing cost through the Publication Assistance 
Program (PAP), Heritage number 09477. 


Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KOG IRO; (613) 269-3211; e-mail: fecook @achilles.net 
Copy Editor: Elizabeth Morton 

Business Manager: William J. Cody, P.O. Box 35069, Westgate P.O. Ottawa, Canada KIZ 1A2; (613) 759-1374 
Book Review Editor: Roy John, 2193 Emard Crescent, Ottawa, Ontario K1J 6K5, e-mail: roy.john @pwgsc.gc.ca 


Associate Editors: Robert R. Anderson Paul M. Catling David Nagorsen 
Charles D. Bird Brian W. Coad Donald F. McAlpine 
Robert R. Campbell Anthony J. Erskine William O. Pruitt, Jr. 


Chairman, Publications Committee: Ronald E. Bedford 


All manuscripts intended for publication except Book Reviews should be addressed to the Editor and sent by postal 
mail. Book-review correspondence should be sent by e-mail or postal mail to Roy John, Book-review Editor. 


Subscriptions and Membership 

Subscription rates for individuals are $28 per calendar year. Libraries and other institutions may subscribe at the rate of $45 per 
year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $28 (individual) $30 (family) $50 (sustaining) and 
$500 (life) includes a subscription to The Canadian Field-Naturalist. All foreign subscribers and members (including USA) must 
add an additional $5.00 to cover postage. The club regional journal, Trail & Landscape, covers the Ottawa District and Local Club 
events. It is mailed to Ottawa area members, and available to those outside Ottawa on request. It is available to Libraries at $28 per 
year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be mailed to: 
The Ottawa Field-Naturalists Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. Canada Post Publications Mail 
Agreement number 40012317. Return Postage Guaranteed. Date of this issue: January—March 2004 (December 2004). 


Cover: A male Northern True Katydid, Prerophylla camellifolia Ottawa, from location 3 Figure 1 in Darbyshire pages 124-126. 
This is the same individual as in Figure 2, page 125. Photographed by Stephen J. Darbyshire. 


MCZ 
LIBRARY 
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FIELD-NATURALIST 


Volume 118 


2004 


Volume 120 
The Ottawa Field-Naturalists’ Club Transactions 


THE OTTAWA FIELD-NATURALISTS’ CLUB 


OTTAWA CANADA 


ate 


The Canadian Field-Naturalist 


Volume 118, Number | January—March 2004 


Origins and History of The Ottawa Field-Naturalists’ Club 


DANIEL F. BRUNTON 


216 Lincoln Heights Road, Ottawa, Ontario K2B 8A8 Canada; e-mail: dbruntonn211@rogers.com 


Brunton, Daniel F. 2004. Origins and history of The Ottawa Field-Naturalists’ Club. Canadian Field-Naturalist 118(1): 1-38. 


The Ottawa Field-Naturalists’ Club (OFNC) represents an unbroken chain of organized, non-governmental natural history 
investigation and education dating back to the early days of the city of Ottawa itself. The Club originated in 1863 with the 
formation of the Ottawa Natural History Society which became the Natural History branch of the Ottawa Literary and Scientific 
Society in 1870, from which the OFNC formally separated in March 1879. Since that time, it has grown into Canada’s oldest 
and largest regional natural history organization and has produced a diverse and internationally recognized publication 
program. Since 1880 The Canadian Field-Naturalist and its predecessors have constituted the scientific core of the OFNC’s 
publication program, with Trail & Landscape being an important Ottawa Valley publication since the late 1960s. The 
importance of both publications to the growth and health of the organization is reflected in the major surges in Club 
membership experienced when each of these publications was established. The focus of membership activities has changed 
over the history of the OFNC, with enlightened natural resource management, then original scientific research and local 
exploration directing energies in the early decades. By the early years of the 20" century the publications program become 
the raison d’etre of the Club, almost to the exclusion of local field activities. A renewed interest in field discovery and the 
growth of conservation awareness in the 1960s, however, rekindled local activities and re-established the balance which has 
sustained the organization throughout its history. Natural environment education has remained a critical theme within OFNC 
programs and activities. Over and above inspiring the professional careers and private interests of thousands of individuals 
for more than a century, the OFNC has had an important and lasting impact on the conservation of natural environment 
features and landscapes in Canada and North America. 


Key Words: Ottawa Field-Naturalists’ Club; Ottawa Natural History Society; The Canadian Field-Naturalist; Trail & Landscape. 


Canada was a mere 12 years old in 1879 when 34 
members of the Ottawa Literary and Scientific Society 
(OLSS) gathered together on an early spring evening 
to discuss a growing problem within Ottawa’s fledg- 
ling naturalist community. Most were full of Victorian 
enthusiasm for discovery and intellectual advance- 
ment and were convinced of the limitless potential of 
their new country. These young men — and they were 
all men, mostly in their 20s or 30s — were frustrated 
by what they saw as an ineffective, moribund OLSS 
Natural History Branch which did not serve their needs. 
These impatient “young Turks” wanted to actually do 
things, to get out into the countryside of the Ottawa 
Valley to explore and discover its natural wonders. And 
with true Victorian missionary zeal, they wanted to 
share these revelations amongst themselves and with 
the larger Canadian — even international — commu- 
nity. All of this in the name of personal intellectual 
development as well as the chance to advance the 
scientific and applied benefits of such knowledge. That 
was heady, revolutionary stuff in the staid, conserva- 
tive Ottawa of March 1879 when how close one was to 
Rideau Hall — the literal and figurative operational base 


of the Governor General and his politically and socially 
powerful entourage — dictated more about one’s status 
and options than personal wealth or political position 
(Gwyn 1984). 

The events which unfolded that cool spring evening 
in the OLSS rooms perched above the muddy streets of 
Ottawa launched careers, changed government policy, 
protected tens of thousands of hectares of Canadian 
natural landscape, produced internationally recognized 
and significant scientific publications, made huge con- 
tributions to our understanding and appreciation of 
North American natural sciences, and enriched thou- 
sands upon thousands of lives. Oh yes ... and initiated 
what has become the largest and oldest regional natu- 
ralist organization in Canada’s history, The Ottawa 
Field-Naturalists’ Club (OFNC). 


The Pioneers (1840s-1863) 

Although the formal beginning of The Ottawa Field- 
Naturalists’ Club in 1879 was 125 years ago, natural- 
ists’ organizations in the Capital actually pre-date Can- 
ada itself (Brault 1946; Dore 1968; Taylor 1986). Prior 
to Confederation, Ottawa (and Bytown before it) was 


D THE CANADIAN FIELD-NATURALIST 


a rude little lumber town characterized more by saw- 
dust, beer and brawls than by intellectual achievement. 
The only adult education or research institution pre- 
sent in those early days was the Mechanics Institute, 
a charitable organization initiated in 1847 as something 
akin to a continuing education facility and library for 
working men. There were no such things as “night 
school” or public libraries, let alone publicly accessi- 
ble research organizations. An informal group known 
as the Silurian Society interested in geological (and 
mining?) issues was also reported to be active in the 
1850s. “Active” may be a misnomer, since they left 
little reference of their doings, other than to suggest 
that their meetings were held “in the City of Ottawa” 
(Anonymous 1854). 

The only natural environment research being under- 
taken in the Ottawa Valley before the 1860s was by 
three highly active individuals. Edward Van Cortlandt 
(1805-1875) was the most socially prominent of these. 
He was one of the first doctors in Bytown, arriving in 
1832 to attend to the military personnel stationed on 
Barracks Hill (now Parliament Hill). He developed an 
extensive private museum of curiosities and artifacts in 
the 1840s, liberally mixing archaeological specimens 
found at aboriginal sites along the Ottawa River with 
natural items dug up, collected, and/or shot in the 
vicinity of the town. He was, by all accounts, a remark- 
ably energetic man who used his high social standing 
to influence local business leaders in natural resource- 
oriented concerns (Moffatt 1986). 

Another dynamo was Elkanah Billings (1820-1876) 
(Figure 1), second son of one of Ottawa’s first pio- 
neer families. Billings was passionately interested in 
natural history in general and geology/paleontology 
in particular. At various times he was a newspaper pub- 
lisher (The Bytown Citizen, forerunner of The Ottawa 
Citizen), a lawyer and finally, Canada’s first profes- 
sional paleontologist. Indeed, this latter science was his 
true calling and he came to be known as “the father 
of Canadian paleontology” (Whiteaves 1876; Clarke 
1971). He presumably was involved in the aforemen- 
tioned Silurian Society, but no direct evidence of that 
was found. The Bytown Citizen was full of natural his- 
tory items during Billings’s tenure (1852-1856), mostly 
representing accounts of his own observations or text 
reprinted from European or American publications. 
These publication activities led directly to his produc- 
tion in Ottawa of Ontario’s first natural science journal, 
The Canadian Naturalist and Geologist, in February 
1856 (Figure 2). Delightful and insightful articles in 
that first volume such as “On the species of woodpeck- 
ers observed in the vicinity of Ottawa” (Billings 1856) 
were based on his extensive travels in the Ottawa 
Valley. They demonstrated both excellent powers of 
observation and a keen appreciation of the importance 
of documenting the appearance and constitution of 
original landscape conditions. His move to Montreal 
later that year to join Sir William Logan at the Cana- 


Vol. 118 


FiGure |. Elkanah Billings. Billings was born in 1820, became 
one of the earliest naturalists in Ottawa, and pub- 
lished the first journal on natural history in Ontario in 
1856 (see Figure 2). He moved to Montreal later that 
year to become the first palaeontologist of the Geo- 
logical Survey of Canada, and the initial curator of its 
museum. (reproduction of the OFNC-commissioned 
portrait, from The Ottawa Naturalist, February 1901). 


dian Geological Survey, however, permanently ended 
his Ottawa connection (Zaslow 1975). 

Elkanah Billings’ older brother, Braddish Billings Jr. 
(1819-1871), completes the trio of Ottawa’s pioneer 
resident naturalists. Braddish was a keen botanist and 
used his position as chief clerk on the Prescott & Otta- 
wa Railway to gain access to a wide variety of habitats 
across eastern Ontario. Although he published nothing 
during this time and relatively little even later, in 1868 
he did produce the first list of vascular plants for the 
city of Ottawa. It was a superb effort for its day, chron- 
icling over 400 species that he found within close 
proximity to the City in 1866 (Dore 1968). Billings 
was seen as a distinguished figure in natural history 
investigations in the Ottawa Valley in the 1860s; like 
his younger brother Elkanah, Braddish was widely 
consulted by natural science researchers elsewhere in 
Canada (Dore 1968). 

The days of exclusively private investigation of the 
Ottawa Valley natural environment ended in the early 


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Waturalist «Grnlogist, & 


By E. BILLINGS, Barrister at Law, 


OTTAWA, CANADA WEST. 


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: UBLISHED AT THE OFFICE or tue OTTAWA CITIZEN, OTTAWA. 


FIGURE 2. The Canadian Naturalist and Geologist, first pub- 
lished in 1856 at the office of The Ottawa Citizen by 
Elkanah Billings, and later continued in Montreal. 


1860s with the transfer of the seat of government of the 
colony of Canada to Ottawa, heralded by the start of 
construction of the new Parliament Buildings in 1860 
(Eggleston 1961). This represented a mega-project with 
huge economic benefits to local businesses. Similarly, 
the 1863 transfer of some 300 bright, educated, and rel- 
atively financially secure members of the civil service 
from the old capital represented both economic and 
social opportunities for the city and the Ottawa Valley. 
Confederation-era city business and social leaders 
quickly embraced the idea of broadening and deepen- 
ing the intellectual resources of the community. In 
furtherance of this, they encouraged the development 
of various clubs and societies to enhance the prestige 
and intellectual capacity befitting the new capital (Brault 
1946; Gwyn 1984). 

It is useful to consider how really rough and ready 
Ottawa and the larger world were in 1863. The Arctic 
was still largely unknown to Europeans and North 
Americans alike but was being charted with great speed 


BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 3 


by the continuing search for the ill-fated Third Frank- 
lin Expedition. The United States Civil War and the 
accompanying ferocious slavery debate raged uncom- 
fortably close to the south, as did intense arguments in 
the Old World regarding the newly published (1859) 
“heresies” of Charles Darwin’s On the origin of spe- 
cies. Closer to home, Ottawa was a bustling, rapidly 
growing city of approximately 15 000 people which 
boasted but a single operating sewer line along Welling- 
ton Street in front of the new Parliament Buildings. The 
first railway train had puffed into town fewer than ten 
years earlier and a municipal drinking-water system 
was still 12 years off (Brault 1946; Eggleston 1961). 
Ottawa was very much straddling the line between 
pioneer lumber town and developing political centre. 


The Ottawa Natural History Society (1863- 
1869) 

It is against this backdrop that a veritable “Who’s- 
who” of the new business and professional elite of 
Confederation-era Ottawa (notably including both the 
aforementioned Edward Van Cortlandt and Braddish 
Billings Jr.) assembled on 3 October 1863 to form the 
Ottawa Natural History Society (ONHS). These gentle- 
men — and the ONHS constitution made it clear that 
membership was open only to gentlemen — were very 
economically oriented. The original minute books 
maintained in the OFNC collection at Library and 
Archives Canada (LAC) records that they were “ ... 
desireous to develope [sic] the Natural History of the 
Ottawa and general resources of the surrounding coun- 
try” (LAC OFNC Collection, 3 October 1863). Explor- 
ing and researching the natural sciences of the Ottawa 
Valley were all well and good but these gentlemen — 
at least initially — wanted to see a profit result from it. 

An important aspect of the growth of such endeay- 
ours was the potential participation of the relatively 
large body of newly arrived civil servants (Brault 1946). 
Activity within quasi-professional associations (it’s 
called “networking” today) was a very important un- 
official avenue for professional advancement within the 
small professional community of public servants in the 
Capital at this time (Gwyn 1984). The ONHS offered 
such an outlet and avenue for civil servants interested 
in natural resources and natural environment issues. 
A definite pecking order mirroring that of their profes- 
sional relationships was soon evident amongst found- 
ing ONHS members who were also senior civil ser- 
vants. When powerful Finance Department Deputy 
Minister John Langton (1808-1894) was ONHS Presi- 
dent in the late 1860s, for example, his ambitious sub- 
ordinate, Interior Department Deputy Minister Edmund 
Meredith (1817-1898), was conspicuously involved in 
the organization, but was careful not to hold higher 
office nor to publicly disagree with Langton (Gwyn 
1984). The social/political significance of the organi- 
zation, over and above the fact that 77 men paid the 
$1.00 membership fee for 1863/1864, can also be meas- 


4 THE CANADIAN FIELD-NATURALIST Vol. 118 


ured by the prominent involvement of individuals such 
as Sir James A. Grant (1831-1920), the Governor Gen- 


eral’s personal physician (Travill 1988) (LAC OFNC Sls = = 
Collection, 3 October 1863). iD = oben re ois 
The ONHS members chose widely appreciated and £ g 2 a 2 z si cs 6 
politically-neutral Braddish Billings Jr. as their first 2) 5 E ee = mm O68 S 
president, although he was not very active in subse- S A lai = = = a Ss ice 
quent Society affairs (LAC OFNC Collection, 3 Oc- 
tober 1863 — 28 October 1864). The Council (Board 
of Directors) soon established a schedule for regular = 
meetings and made arrangements for the development s = 
of “The Cabinet’, a series of cases displaying natural ARE Aer mes = E 
: 3 E 3 ‘oO oo av 2 >) 
history specimens which were to be held in the rooms eg Weipa) ds se 2 
they rented at the Mechanics Institute building on ea Bg Bi = & mm Ss 
Sparks Street. Van Cortlandt was elected museum coho areas Re) ead Teal 
curator for the Society and remained in that position 
throughout the life of the organization. s 
Although precise records are sketchy, records at a os = 
Library and Archives Canada indicate that the Society eS ees Saas 
: Xe) os oY & o 
met regularly for lectures and field trips. The last C2 Milfsseehy ssi Sho 2 
Saturday of each month was fixed for the latter (LAC Ss 8 ae S = = OS 
OFNC Collection, 28 April 1865), during which mem- chr een ee areal i age bad ifs teal 
bers actively searched for natural history specimens for 
The Cabinet. Indeed, this seemed to be a major focus s 
of the organization (Dore 1968). In October 1869, only wee & 
months before the organization’s amalgamation with ur 2 ge gel es 5 
the Mechanics Institute, Braddish Billings offered his ca) = = Se Ss a 
herbarium to the ONHS for $60 (LAC OFNC Collec- Ra ES Es & = & A s 
tion, 1 October 1869). Although no supporting motion Co Zagtesie tes te ee} (e\ al 
for such a purchase is noted, Billings’s collection did 
end up there (it may have been deposited after his 3 
death), and was examined in the OLSS museum in the 5 oa thie & 
1880s (Fletcher 1888). Unfortunately, the OLSS herbar- e [2+ Bee Fs 5 
ium, including the Billings’s specimens, disappeared 2 bere seee 2 
sometime in the early 20" century (Dore 1968). g Ali olepese Haya daeiply 
The ONHS seems to have been most active between oe ea PA (as esis ea eh teal 
1865 and 1867 and to have achieved considerable ic} 
standing in the community. In 1866 a group met with S D es 
federal Minister Thomas Darcy McGee to discuss a oe in 2 
possible ONHS — Mechanics Institute exhibit in the = 8 eae te % : 
1867 Paris Exhibition (LAC OFNC Collection, 26 SS ie a Be a2 S 
April 1866). Their paths had crossed before, in Que- >a Ie a ae os 3 
bec City in 1862-1863, McGee had lectured to the Ss1% BS =S <Os 3% 
Quebec Literary and Historical Society along with a of 
both E. A. Meredith and John Langton. And while the aS 
subject of his speech was not recorded, the Governor = = : ‘2 % 3 = 
General apparently spoke to the Society shortly there- é 3 a a 8 aie) ae = S 
after (LAC OFNC Collection, 25 May 1866). Within PSS |S 252 88) vols 
a month of Confederation an ONHS delegation led by 5 2 SS a § iz § gx Salles 
President N. B. Webster lobbied Prime Minister John sE IS c = a ss = HAs z = 
A. Macdonald to have the Geological Survey Museum a. || it Ts Sa es 
moved to Ottawa from Montreal (LAC OFNC Collec- g 3 5 
tion, 26 July 1867) — possibly the first such initiative in aie, i = 
the lengthy campaign to effect such a transfer (Zaslow 3Z & 5 BS, 
1975). Similarly, early concerns for habitat protection 2 5 2 ¥ = 3 
and landscape conservation were discussed amongst the a Q co pee tn i g SS 
Society’s influential membership through presentations ae 8 2 gs 2 ay S 5) 
like Vice-President Thomas Austin’s lecture “The UI- a & i ae 5 3 ESs|2 
ae pa ea el (455) || 


2004 


terior Effects of Clearing off the Forests and Draining 
the Country” (LAC OFNC Collection, 26 April 1866). 

While The Cabinet may not have survived, the Soci- 
ety left a more lasting (albeit, modest) record through 
its publications program. It commenced in 1867 with a 
pamphlet on the possible economic uses of Milkweed 
(Asclepias syriaca) by Alexander Kirkwood (1823- 
1901) who, with the timely promotional assistance of 
ONHS member and OFNC founder Henry B. Small 
Sr. (1832-1919), would later be the driving force be- 
hind the establishment of Ontario’s Algonquin Prov- 
incial Park (Killan 1993). The ONHS published 250 
copies of Kirkwood’s lecture (Kirkwood 1867). Ap- 
proval for the production of 250 copies of a lecture by 
Van Cortlandt entitled “Native Compounds and Metal- 
lurgy of Iron” was given by the Council the previous 
year but there is no evidence that a publication actually 
resulted (LAC OFNC Collection, 28 December 1866). 
After a year of discussion and buoyed by the success of 
the Kirkwood pamphlet, however, an ONHS Trans- 
actions series was initiated. It was short lived, produc- 
ing only three numbers. ! 

Faced with rising financial pressures (including the 
costs of renting rooms) and the opportunity to formally 
combine resources (including a provincial operating 
grant), the ONHS decided in the spring of 1869 to 
merge with the Mechanics Institute (LAC OFNC Col- 
lection, 16 April 1869). The minutes books describe 
surprisingly little of the ensuing discussion, but the 
Society did merge with the Institute to form the Ottawa 
Literary and Scientific Society (OLSS) on 24 December 
1869 (LAC OFNC Collection, 30 December 1869). The 
ONHS Cabinet became the basis for the OLSS muse- 
um, whose collections disappeared with the dissolution 
of the Society in 1906-1907. Unpublished biological 
data were apparently also maintained in the Museum, 
however, as indicated by later reference (containing 
several errors in fact) being made to “lists published 
by the Ottawa Natural History Society of 1859-63” 
(Small 1883). 

Although active for only a decade, the ONHS was 
critical in the awakening of interest in the organized 
and documented investigation of natural sciences in the 
Ottawa Valley. It bridged the gap from hit-and-miss 
personal interest to organized and semi-professional in- 
vestigation. It also initiated the concern for and partici- 
pation in larger issues of national science policy, re- 
search priorities, and natural environment conservation 
which continue within the Ottawa naturalist commu- 
nity to the present day. 


The Ottawa Literary and Scientific Society 
and OFNC formation (1870-1879) 

The combination of the ONHS and the Mechanics 
Institute appears to have been seamless, with the mem- 
bers of the ONHS effectively becoming the Natural 
History Branch of the newly created OLSS. Little 
documentation was found, however, concerning the re- 
sulting organization in general and the activities of its 


BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 5 


Natural History Branch in particular. Nonetheless, the 
OLSS was a socially and culturally significant orga- 
nization in Victorian Ottawa by the end of the 1870s, 
supporting a regular lecture series, a public reading 
room and a lending library of 1100 volumes (LAC 
OFNC Collection, OLSS brochure). Despite the larg- 
er city population, better local research resources and 
the existence of a potential sponsoring organization, 
however, there are few indications of more than indi- 
vidual efforts in natural science investigations in the 
Ottawa Valley. At the least, the momentum of the 
1860s seems to have been stymied within the larger, 
predominantly culturally-oriented OLSS. The stage was 
set for a final transformation of the ineffective OLSS 
Natural History Branch into a new and more produc- 
tive structure ... an independent naturalist group. 

If the organization that morphed into the OFNC 
actually began in October 1863, the final stage of its 
transformation was achieved with the cordial separa- 
tion of the Natural History Branch from the main body 
of the OLSS in March 1879. It appears to have been 
a classic example of a group of activists becoming 
dissatisfied with the pace and effectiveness of a well- 
established group and deciding that the only way to 
achieve more contemporary goals was to strike out on 
their own. The fact that most of the prime movers and 
shakers in the new organization were established mem- 
bers of the old OLSS and remained members for years 
thereafter, indicates that they retained faith with the 
original organization as a valuable forum for intellec- 
tual development and social debate. Nonetheless, they 
wanted to see more activity and greater opportunities 
being provided for natural environment investigations 
(Harrington 1909). 

Once again, national and world affairs played an 
important role in the development of naturalist organi- 
zations in the Ottawa Valley. Canada was experiencing 
the early years of a serious economic downturn that 
lasted into the early 1890s (Eggleston 1961). At the 
same time, the promise of this new country and the 
political, social and economic difficulties of the Old 
World were encouraging huge numbers of young, rela- 
tively well-educated and mobile immigrants to enter 
Canada; some stayed and some moved on in this time 
of social and economic upheaval. The decade follow- 
ing the founding of the OFNC, for example, saw the 
highest number of both Canadian immigrants and emi- 
grants of any time in the 19" century (Lingard 1967). 


Young Turks and Rebels (1878-1879) 

The group which gathered in March 1879 to con- 
sider a new natural history organization in Ottawa re- 
flected its times. This included a mix of Ottawa vet- 
erans enthusiastic about the future, as well as young, 
newly-established civil servants, and British immigrants 
full of imperial fervour. They seem to have been imbued 
with Victorian optimism about the importance and 
strength of ideas, a missionary zeal for discovery and 
the sharing of knowledge, and a strong desire to con- 


6 THE CANADIAN FIELD-NATURALIST 


tribute to their community (LAC OFNC Collection, 
25 March 1879-15 December 1879). Some stayed on 
to become important participants in the exploration of 
the Ottawa Valley natural environment while others fol- 
lowed different paths elsewhere in Canada and beyond. 

The founding meeting was called by James Fletcher 
(1852-1908), OLSS museum curator since 1878 and 
an up-and-coming entomologist and botanist (Figure 
4). Fletcher had emigrated from England in 1874, mov- 
ing to Ottawa in 1875 to work as a clerk in the Bank of 
British North America and joining the Parliamentary 
Library staff as an accounting clerk in 1876 (Cody et 
al., 1986). He was a gregarious, personable, and phys- 
ically active man who quickly became deeply involved 
in sporting and social activities. He was, for example, 
a keen snowshoe racer and a founder and principal 
player with the Ottawa Football Club, a rugby team 
which later evolved into the Ottawa Rough Riders Can- 
adian Football League team (Harrington 1909). He was 
also a pillar of the Anglican Church community in 
Ottawa and remained so throughout his life. And, while 
being a young man with shallow Ottawa roots, his so- 
cial profile, and thus influence, improved dramatically 
when he married Eleanor Schreiber (daughter of Sir 
Collingwood Schreiber, Chief Engineer of the Canadian 
Pacific Railway and later Chief Engineer of the Depart- 
ment of Railway and Canals) in 1879. 

Fletcher’s enthusiasm, intelligence, and boundless 
energy quickly became known to the small group of 
active field naturalists already resident in Ottawa and he 
was eagerly sought out (Whyte 1909). Chief amongst 
these field associates and life-long friends were Robert 
B. Whyte (1851-1918), an Ottawa-born private busi- 
nessman and passionate horticulturalist (Macoun 1918) 
(Figure 5). The other was William (Will) H. Harring- 
ton (1852-1918), a skilled entomologist and botanist 
who moved from Nova Scotia in 1870 and spent his 
entire working career with the federal post office 
department (Gibson 1918). While Whyte is credited 
with the idea of an Ottawa Field-Naturalists’ Club, 
Fletcher is universally credited with being the fellow 


Vol. 118 


who made it happen and saw to it that it became firm- 
ly established (Harrington 1909; Whyte 1909).? 

The dynamic trio of young naturalists were not alone 
in this, and were joined by an eclectic mixture of OLSS 
members who also felt the need for an organization 
focusing exclusively on natural sciences. Many were 
prominent in the building of Ottawa and/or the federal 
civil service of the new country (Brault 1946; Gywn 
1984). These included former ONHS officer and senior 
Post Office Department official William White (1830- 
1911) (Figure 3); lawyer and politician (later, Premier 
of British Columbia) Joseph Martin (1852-1923); for- 
mer ONHS officer, teacher, and Geological Survey of 
Canada (GSC) librarian John Thorburn (1830-1904); 
author and later Deputy Minister of the Interior Henry 
Small Sr., and the OLSS president of the day, promi- 
nent literary patron and Secretary (Deputy-Minister) 
of the Post Office Department, William D. LeSueur 
(1840-1917) (LAC OFNC Collection, 25 March 1879). 

As noted earlier, the social and political prominence 
of members was important to the success of such or- 
ganizations in Victorian Ottawa (Gywn 1984). This 
early naturalist community (Table 2) was small and 
remarkably homogenous, dominated by white, English- 
speaking, Anglo-Saxon protestants who worked, lived 
and even were buried in close proximity. Overwhelm- 
ingly, the children of most early club leaders were 
enrolled in the Ottawa Collegiate Institute (MacMillan 
et al. 1904), and numerous Club officers — including 
almost two dozen former Presidents — are buried at 
Beechwood Cemetery. Beechwood, appropriately 
enough, was also a favoured 19" Century OFNC ex- 
cursion site (Reddoch 1979c). 


Founding Meeting (25 March 1879) 

The special meeting of OLSS Natural History 
Branch members was called to order in the OLSS 
museum at 112 4 Sparks Street by James Fletcher (Fig- 
ure 4) on 25 March 1879. It seems that the politics 
of Victorian Ottawa immediately intervened, however, as 
William White (Figure 3) and R. J. Wicksteed moved 


TABLE 2. Participants at inaugural meeting of the OFNC, 25 March 1879 


(LAC OFNC Collection, 25 March 1879). 


W. P. Anderson L. A. Hamilton Dr. Ross 

E. D. Arnaud W. H. Harrington H. B. Small Sr. 
Prof. George Baptie G. Heron H. B. Small Jr. 
W. R. Billings G. A. D. Jones P. D. Symms 

W. Chesterton E. V. Johnson John Thorburn* 
L. H. Chrysler W. D. LeSueur* H. Watters 

R. A. Davy J. Martin W. White* 
James Fletcher B. Monk R. B. Whyte 

J. M. Greta S. McLaughlin* R. J. Wicksteed* 
J. MacD. Gordon Prof. W.R. Riddell A. P. Wright 


J. A. Guignard 
D. Horsey 


*member of ONHS (1863-1869) 


C. J. Ripley 
P. Robertson 


2004 


FiGurRE 3. Lieutenant-Colonel William White, May 1901. White 
was the first president of the Ottawa Field-Naturalists’ 
Club, and also the Ottawa Horticultural Society. Earlier 
he was president of the Ottawa Athenaeum and Mech- 
anical Institute. Photographer William James Topley, 
Ottawa, archived at the Library and Archives of Can- 
ada, Ottawa: E 81666). 


that fellow ONHS veteran John Thorburn chair the 
meeting. Perhaps to counter this old guard move, new- 
comers Joseph Martin and Henry B. Small Jr. subse- 
quently moved that Fletcher be made Secretary of the 
meeting. The social pecking order apparently satisfied, 
Fletcher then got discussion under way (LAC OFNC 
Collection, 25 March 1879). 

After some debate of whether to stay affiliated with 
the OLSS or to form a separate group, the actual mo- 
tion to establish an independent OFNC was moved 
by Joseph Martin (seconded by R. B. Whyte): “... 
that it is advisable to form a Field Naturalists Club 
for the City of Ottawa and do proceed to organize”. 

It was only after this motion had been passed that 
Fletcher reported to the gentlemen assembled that a 
small group (consisting of at least James Fletcher, Will 
Harrington, R. B. Whyte (Figure 5), Joseph Martin, 
and Henry Small Jr.) had actually held preliminary 
consultations on 11 March 1879 to prepare a proposal 
to the Council of the OLSS — and no doubt, to map 
out strategies. At that preliminary meeting they had 
drafted a motion (Fletcher, seconded by H. B. Small 
Jr.) that in light of “... an Ottawa-Field Naturalists 
Club having been organized in Ottawa...”, this new 


BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 7 


group “... would gladly contribute to the Museum of 
the OL & S Society and thereby revive its present 
dormant condition”. In exchange, they asked that the 
OFNC be allowed to use the OLSS rooms for meetings. 
An additional clause asking that the new organization 
be “under the auspices” of the OLSS was an apparent 
face-saving offer, since the OFNC had virtually noth- 
ing further to do with OLSS programs.’ The resolution 
was passed by the OLSS Council, though there likely 
was little else they could do, being presented with such 
a fait accompli. There may have been a good bit of bluff 
on the part of Fletcher and his associates, however, 
since the Club did not in fact exist on 11 March when 
the motion was drafted! 

If this was not-too-subtle political manipulation by 
the newcomers, the old guard may well have reaped a 
bit of revenge in the election of OFNC officers which 
followed immediately thereafter. William White, long- 
time Ottawa resident, former Mechanics Institute pres- 
ident and ONHS member, was elected President. It is 
not recorded in the Minute Books if the officer positions 
were contested that day, as was often the case in the 
first years of the organization. Likely more important 
than being an ONHS veteran, White was a senior Post 
Office Department official and was socially prominent 
as Lieutenant-Colonel of the Governor General’s Foot 


Ficure 4. James Fletcher in June 1908. He called the founding 
meeting of The Ottawa Field-Naturalists’ Club and was 
the prime architect of its early success. Photogra- 
pher William James Topley, Ottawa, archived at the 
Library and Archives of Canada, Ottawa C96620). 


8 THE CANADIAN FIELD-NATURALIST 


Guards (Shutt 1912; Anonymous 1929). His election, 
however, may very well have also been the wish of the 
newcomers since White’s Rideau Hall connections 
made the important task of securing the agreement of 
the Governor General to serve as Club Patron that 
much easier. 

The minutes of the founding meeting (LAC OFNC 
Collection, 25 March 1879) are very sparse in detail. 
As Secretary, James Fletcher would have produced a 
hand-written record of events. The preserved minutes 
are in the handwriting of R. B. Whyte, however, indi- 
cating that Fletcher’s record of events was rewritten. 
The development of this simplified record could per- 
haps have allowed for the tactful omission of politically 
delicate matters such as contested positions and/or 
heated exchanges. 

In any event, James Fletcher was elected First Vice- 
President and the Council and Officer positions were 
filled by other young rebels (R. B. Whyte, Henry B. 
Small Jr., W. H. Harrington, and Joseph Martin) or 
neutral figures (W. R. Riddell and J. A. Guignard) 
(Appendix 1). Only one Council position was filled 
by a White associate, William P. Anderson (1852- 
1927), who also later became Lieutentant-Colonel of 
the Governor General’s Foot Guards (Small 1929). 
As it turned out, however, Fletcher essentially ran the 
affairs of the OFNC in 1879 and 1880, with White 
playing a very low-key role. 


Earliest Days (1879 — 1880) 

The first Council meeting of the new Club was held 
under the chairmanship of William White on the after- 
noon of 3 April 1879 in the Museum of the OLSS 
(LAC OFNC Collection, 3 April 1879). It was fol- 
lowed almost immediately by another, longer session 
at the home of William Anderson on 5 April 1879 to 
prepare details for a formal members’ General Meet- 
ing the following Tuesday (8 April 1879) and to accept 
the resignation from the Council and from the Club of 
J. A. Guignard. No reasons were given for the speedy 
and rather dramatic resignation of Guignard but des- 
pite being James Fletcher’s assistant at the Experi- 
mental Farm from 1891 onward, he did not ever re- 
join the Club. He did, however, function as Acting 
Editor of The Ottawa Naturalist for several months 
in 1903 (Brunton 1986a). 

The Ottawa Field-Naturalists’ Club first met public- 
ly on 8 April 1879 under the chairmanship of James 
Fletcher with about 25 members in attendance in the 
Museum of the OLSS. Operational rules for the Club 
were described, a Corresponding Members (Honorary 
Member) designation was identified, an active excur- 
sion program was laid out, and a membership fee 
(SO0¢ per annum) was established. The fee was half 
that established by the ONHS 16 years earlier, presum- 
ably in an effort to keep membership in the OFNC 
more affordable for the general public. And also un- 
like the ONHS, the OFNC specifically identified mem- 


Vol. 118 


bership as being open to both “ladies and gentlemen 
desiring to join the Club” (LAC OFNC Collection, 8 
April 1979).° 

There were 25 OFNC Council and public meetings 
held in 1879/1880, the busy first year of the newly 
independent Club. Most of the public meetings were 
in the form of somewhat informal soirées in which 
lectures were followed by opportunities for debate and 
conversation. Through this year the Council addressed 
numerous details of the developing organization in- 
cluding the arrangement of Vice-regal patronage, the 
establishment of an active excursion program, the 
selection of the first OFNC Corresponding Member 
(John Macoun [1831-1920]) (Brunton and Gummer 
1987), and recognition of the importance of “public 
education” (the first gimmers of conservation action?). 
James Fletcher was everywhere in this, conducting ex- 
cursions, conducting and documenting field research, 
and handling many logistical details. President White 
chaired 12 of the meetings that year, only a few more 
than Fletcher, who oversaw nine because of the presi- 
dent’s frequent absence (LAC OFNC Collection, 1879- 
1880). 

The establishment of various scientific working com- 
mittees (botany, ornithology, geology, etc.) generated 
a great deal of interest and field activity within the 
Ottawa District®, resulting in considerable natural en- 
vironment information being gathered. Membership 
grew to almost 100. The influence of Lieutentant- 
Colonel White and others secured the patronage of 
the Governor General and made OFNC membership 
socially desirable. This was aided by the membership 
within the first year of prominent individuals like the 
previously-mentioned Governor General’s personal 
physician, Sir James Grant; the founder of the Central 
Experimental Farm research institution, Sir William 
Saunders (1836-1914) (Anstey 1988; Cody et al., 
1986); and prominent scientist, engineer, and inventor 
Sir Sandford Fleming (1827-1915) (Regehr 1988). 

The lectures delivered at the Soirées were well 
received, leading to requests for their publication for 
the benefit of members and other interested parties 
alike. The March 1880 Council meeting established 
that a transaction of the OFNC’s first year be produced. 
Five hundred copies of that first Transactions were 
published for a cost of $78.43, likely in June 1880 
(Brunton 1986a) (Figure 8). In appreciation of his 
production of the lithographed plates for the publica- 
tion at no cost, the Council granted Club membership 
for the year to J. A. Guignard — the same gentleman 
who had resigned from the Council as well as from 
Club membership, only days after the Club had been 
formed (LAC OFNC Collection, 28 July 1880). This 
was done again in 1882 for additional gratis litho- 
graphic services, so it appears that whatever difficulty 
Guignard had with being on the Council and being a 
voluntary Club member, he was prepared to receive 
and work for the publication. 


—— 


2004 


FicurE 5. Robert B. Whyte, (June 1908), was first secretary- 
treasurer and later president of the Ottawa Field- 
Naturalists’ Club. Photographer William James Topley, 
Ottawa, archived at the Library and Archives of Can- 
ada, Ottawa: C 105521). 


Publication of the Transactions was the first step 
in a renowned publication program that, to date, has 
produced seven annual Transactions of The Ottawa 
Field-Naturalists’ Club, almost 120 volumes of The 
Ottawa Naturalist/The Canadian Field-Naturalist and 
almost 40 volumes of Trail & Landscape. Complete 
sets of The Canadian Field-Naturalist and its prede- 
cessors are rare, however, particularly so in private 
libraries. Likely fewer than a dozen complete sets 
exist in Canada. The Transactions are especially rare 
publications, several of which became unavailable 
within a few years of publication. Only 75 copies of 
Transactions | were still available by early 1884, for 
example (LAC OFNC Collection, 12 March 1884), 
and only “virtually complete sets” were being sold (for 
$24) by 1909 (LAC OFNC Collection, 23 February 
1909). 


The Late Victorians (1880s-1890s) 

The OFNC was a huge hit at the height of the Vic- 
torian era in Ottawa. Aided by improved transportation 
— most particularly, the expansion of railways up the 
Ottawa and Gatineau Valleys — and the continued 
growth of both the federal civil service and the city 
(exploding to 60 000 by 1899 — Brault 1946), huge 


BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 9 


strides were being made in the exploration and under- 
standing of the natural biodiversity of the Ottawa Valley. 
Transfer of the Geological Survey of Canada from 
Montreal to Ottawa in 1880 (Zaslow 1975) and with 
it, many of the top natural scientists in Canada, pro- 
vided a major boost to the OFNC. So too did the 1882 
arrival of John Macoun (Macoun 1922). Through their 
network of professional connections, Macoun and other 
scientists at “The Museum’, as it was known, also 
provided an important link between OFNC members 
and national and international floral and faunal au- 
thorities. Despite becoming increasingly preoccupied 
with professional duties after he became Dominion 
Entomologist and Botanist in 1886 (Cody et al., 1986) 
and being troubled by serious eyesight problems for 
a period of time (LAC OFNC Collection, 17 March 
1885), James Fletcher remained an inspirational and 
hands-on leader within the OFNC throughout the 1880s 
and 1890s. 

Club membership had more than doubled to 254 by 
1899 (Figure 6). This was due to both local and wider- 
scale influences. On the local front, OFNC excursions 
became major social events and major generators of 
additional interest in the organization (Reddoch 1979c). 
These were often huge events, with trains being char- 
tered to take upwards of 300 participants on day-long 
trips. Fortunately, many sites now well within the urban 
core of the National Capital Region were in close to 
original condition then and became prime locations for 
many formal and informal outings. Chief amongst these 
were Lac Leamy, Fairy Lake, and Wychwood (Aylmer) 
in Gatineau, and Rockcliffe Park, Beechwood Ceme- 
tery and the Billings Bridge (Rideau River) area in 
Ottawa. Further afield, the huge Mer Bleue peat bog 
and Casselman to the east, and King Mountain and 
Chelsea to the north offered “exotic” destinations that 
were visited repeatedly (Reddoch 1979c). 

In striking contrast to the early to mid-19" century 
days of the ONHS and its predecessors, these efforts 
were well documented in OFNC publications. The 
annual Transactions, modelled on the Transactions 
of the Manchester Field-Naturalists’ Society (Brunton 
1986a), were produced through much of the 1880s. 
Each Transactions reproduced Club lectures delivered 
during the previous year. More importantly, they includ- 
ed annual reports of various committees highlighting 
the field discoveries of members. The Transactions 
thus provided the first documentation of a variety of 
comprehensive lists of Ottawa area flora and fauna, 
such as birds (White and Scott 1882), vascular plants 
(Fletcher 1880), and fish (Small 1883). Against these, 
individual members could compare and direct their own 
field investigations and discoveries. Fletcher’s serial- 
ized, annotated revision of the 1880 list of vascular flora 
was produced over many years thereafter, providing a 
continuing enumeration of new discoveries and poten- 
tial exploration sites for OFNC members (Boivin and 
Cody 1955). 


10 THE CANADIAN FIELD-NATURALIST 


After seven annual Transactions had been produced, 
there were increasing calls for a more frequently 
appearing publication. This was particularly important 
for the description of new species (LaRocque 1931), 
the first being a new fossil which had been described 
in Transactions 2 by Sir James Grant (Grant 1881). 
At the March 1887 Annual Meeting members agreed 
to initiate a monthly publication series which could 
move beyond the simple publication of Club lectures 
and reports. This was over the objections of W. P. 
Anderson, the current OLSS President, who had com- 
plained the previous fall of OFNC “antagonism” to- 
wards the old Society (see above); he wanted the 
Transactions to stay as they were (LAC OFNC Collec- 
tion, 14 March 1887). 

The first issue of The Ottawa Naturalist was pub- 
lished in April 1887 under the editorship of Will Har- 
rington, who also had headed up the editorial com- 
mittee which produced the last (1886) volume of the 
Transactions (Brunton 1986a). The Ottawa Naturalist 
was a small (21 x 14 cm), slim (16 page) publication 
produced on a shoe-string budget (150 copies at $18.50 
per issue) (LAC OFNC Collection, 10 March 1887). 
Even then, such a cost was considered a financial dif- 
ficulty for the Club (Harrington 1887) but was off-set 
to some degree by the sale of advertising space on the 
back cover. These advertisements included notices to 
such “critical” products and services for the attention 
and consideration of Ottawa naturalists as G. W. 
McCullough’s Anthracite and Bituminous Coal, C. 
Ross & Co.’s “Beautiful selection of Dress Silks being 
offered at 75¢ per yard”, and Miss Harmon’s Board- 
ing and Day School for Young Ladies. 

Not only the advertisements may have met with 
amusement or disapproval in the eyes of some. In 1888 
the always feisty John Macoun expressed the opinion 
that “too much trash was now published”, and that 
“the majority of papers were of little or no value ...”; 
that the accounts of Excursions were “all but worth- 
less”. It is not recorded what Editor Will Harrington 
might have thought upon hearing this (he undoubtedly 
was present), but OFNC president R. B. Whyte was 
not amused by Macoun’s intemperance. Whyte stated 
that he “ ... wished to record his expression as direct- 
ly opposite [Macoun’s] and thought the publication 
was of much interest to the members” (LAC OFNC 
Collection, 9 March 1888). 

Regardless of the reservations of people such as 
William Anderson and John Macoun, The Ottawa 
Naturalist established a new standard for the timely 
presentation of technically sound, original scientific 
information which has continued seamlessly to the 
present day. Its launch, however, was also far from the 
last time controversy and disagreement would charac- 
terize discussion and debate about OFNC publications! 

Both the growth in local capacity for natural envi- 
ronment research and the evolving vision of the Club’s 


Vol. 118 


role are indicated by Council’s November 1887 rejec- 
tion of a gift of biological specimens from Corres- 
ponding Member and American entomologist Henry 
Edwards (1830-1891) (Fletcher 1891). The Council 
(with both John Macoun and James Fletcher present) 
suggested that the potential donor should offer the 
plant specimens to the herbarium of the Central Exper- 
imental Farm (DAO) which had recently been started 
by Fletcher and is now the largest collection in Canada 
(Rothfels 2003); they further suggested that Edwards’s 
insect specimens be offered to the Geological Survey 
Museum (LAC OFNC Collection, 18 November 
1887). There was no discussion of a donation to the 
OLSS collection, nor of initiating an OFNC museum 
or cabinet. Neither, it appears, was deemed appropri- 
ate in light of the existence of these growing, pro- 
fessionally based research collections. 

By the end of the 19" century, the personality of 
the Club had also evolved considerably. The OFNC 
was now administratively well established and boasted 
a program of regular meetings and excursions. It en- 
joyed a strong reputation for community involvement, 
produced a steady stream of field-based scientific find- 
ings, and even demonstrated a measure of political 
prominence with Laurier government cabinet minis- 
ters and future provincial Lieutenant-Governors as 
members. The parent organization from which it had 
split off, on the other hand, was in serious decline and 
only a few years from dissolution. The young turks who 
had rescued the OFNC from the moribund OLSS 
Natural History Branch were middle aged and well 
established now, a number having become respected 
senior scientists and policy makers. The Ottawa Nat- 
uralist remained a monthly publication but had grown 
in size and enjoyed much improved printing quality. 
Articles often now were accompanied by illustrations, 
these sometimes being photographic. The Ottawa Nat- 
uralist was gaining a wider readership too, with articles 
addressing subjects considerably further afield than 
the Ottawa Valley, such as the review of the status of 
bird species in King’s County, Nova Scotia (Tufts 
1898; 1899a; 1899b). 

But portents of future conflicts and a growing diver- 
gence of visions could be sensed. Founder and Past- 
president R. B. Whyte, for example, complained about 
the Nova Scotia bird articles. He felt The Ottawa Nat- 
uralist should not be publishing material so far removed 
from the Ottawa area. Botanist James M. Macoun 
(1862-1920) and geologist Henry Ami (1858-1931) 
disagreed, suggesting that the Club publication fulfilled 
an important role here. Other new members support- 
ed Macoun and Ami and called for a greater number 
of rigorously scientific, more broadly based articles to 
be included in the publication (LAC OFNC Collection, 
14 March 1899). 

In other words, a new group of younger, more vig- 
orous, field-oriented naturalists — including a number 


2004 BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 1] 


Number of Members 


assis ctor ieee 


Ficure 6. Ottawa Field-Naturalists’ Club membership trend, 1879/1880 to 1974. Membership lists until 1971 (published in 
The Canadian Field-Naturalist until the early 1950s and/or reported in published Annual Reports) included all 
addressees to which The Canadian Field-Naturalist was sent. After 1971, however, the membership totals exclude 
non-voting subscribers and thus are no longer directly comparable to earlier numbers. Where gaps on the membership 
data occurred, such as when no lists or Annual Reports were published in some years in the 1920s, an intermediate 
estimate was entered. 


FiGureE 7. A geological field trip of The Ottawa Field-Naturalists’ Club along a railway rock cut north of Chelsea Grove in the 
Gatineau region of Quebec, north of Ottawa. The area’s oldest and youngest deposits are in direct contact here. The trip 
leader was Dr. Henry Ami. From Trail & Landscape 13(3): 94; prepared from a lantern slide original by Dr. Ami, 
now in the Library and Archives of Canada, Ottawa. 


12 THE CANADIAN FIELD-NATURALIST 


4879. 


OTTAWA 


Pe ‘ 


FIELD- NATURALISTS’ CLUB. | 


TRANSACTIONS No. 14. 


OTTAWA, CANADA: 
CITIZEN PRINTING AND PUBLISHING COMPANY, SPARKS STREET. 


1880. 


FiGureE 8. The initial issue of The Ottawa Field-Naturalists’ 
Club Transactions, the initial annual publication series 
of The Ottawa Field-Naturalists’ Club. 


of the first wave of trained professional biologists to 
work in the federal government — were becoming 
prominent in the organization, changing things, and 
arguing for a more national perspective. This was not 
entirely to the liking of at least some of the long- 
established Club leadership. The situation must have 
had ironic echos for the elders of the Club like Whyte, 
Harrington, Small, and Fletcher, for in the days 
leading up to the founding of the Club they were the 
“young turks” eager for change! 


Memorials and the Great War (1900-1915) 

Despite the economic trials of much of the late Vic- 
torian era, it was a positive period of establishment, 
growth and achievement for the OFNC. By the turn of 
the 20" century, it had become one of the largest and 
most prominent such organizations in Canada. Events 
of the next decade and a half would sorely test that 
status, however. 

Symbolic of the coming of age of the OFNC and 
the end of its establishment period was the death of 


Vol. 118 


Queen Victoria in 1901. The OFNC pioneers, after all, 
were either originally emigrants from Great Britain or 
were locally-born in colonial times. The black-edge, 
memorial issue of The Ottawa Naturalist in February 
1901 (Figure 9) expressed in its frontispiece tribute 
the heart-felt distress at the passing of an era: “In com- 
mon with all the sorrowing subjects of His Imperial 
Majesty King Edward the Seventh the members of 
The Ottawa Field-Naturalists’ Club desire to record 
their deep sense of sorrow and loss at the demise of 
their beloved Sovereign Lady, Queen Victoria, during 
whose glorious reign of sixty-four years, scientific work 
and original research, such as our Club aims to accom- 
plish, have received unprecedented impetus”. Quite so! 

But Victoria was not the only recent loss mourned 
by the Club in the new days of the new century. In the 
Victoria Memorial edition, in fact, President Henry 
Ami laments the February 1899 “early demise of our 
friend and fellow member, the sweet poet of Ottawa, 
Archibald Lampman ....[whose] ardent love of Nature 
and all she teaches in lake, forest, in autumn, in winter, 
in sorrow, in comfort, led him into those numberless 
nooks and sequestered spots which enchant the eye, 
please the mind and entrance the soul” (Ami 1901). 

Just to be sure that the readers not think the Club 
had become overwhelmed by anthropocentric concerns, 
Ami went on in his address to eulogize a major 
scientific personality ... the aforementioned Elkanah 
Billings who had played such an important role in 
the founding of scientific natural environment investi- 
gation in the Ottawa Valley in the first place. Ami 
further announced that the Club has commissioned a 
portrait of the late Mr. Billings and was donating it to 
the Geological Survey Museum (Figure 1).’ Shortly 
after Ami’s address, the Council established a com- 
mittee to co-ordinate with “different societies affiliated 
with the Royal Society of Canada” lobbying of the 
federal government for the construction of a National 
Museum (LAC OFNC Collection, 13 February 1900). 
Was this the genesis of the undertaking which resulted 
in the completion and occupation of the magnificent 
Victoria Memorial Museum building in 1910-1911? 
In any event, the Club officers were clearly aiming for 
a balance between the scientific investigation of the 
natural environment and the encouragement/enhance- 
ment of public awareness of the importance of such 
concerns. 

A growing OFNC interest in and emphasis on en- 
couraging public awareness and appreciation of natural 
sciences is shown by the active promotion of involve- 
ment by the students and staff of the Ottawa Normal 
School (teachers college) during the early years of the 
20" century. This was the dawning of “environmental 
education” in North America and the Club seemed 
enthusiastically involved, particularly through the 
school. At least five members of the OFNC Council 
worked at the school in this time, including Vice- 
Principal S. B. Sinclair. Sinclair served in various Club 


2004 


capacities during this period, including OFNC 1905- 
1906 President (Appendix 1). 

Not everyone shared this positive view of environ- 
mental education within the Normal School and Ottawa 
public schools. A lengthy debate was held amongst 
key Club members early in 1905 on the difficulties and 
obstacles of this undertaking. In his characteristically 
direct way John Macoun declared “it to be his con- 
viction that Nature Study would soon be consigned 
to the limbo of exploded educational fads” (Attwood 
1905). Nonetheless, the Club remained formally in- 
volved in such school programs for years thereafter. 

OFNC excursions also remained popular public 
events, with over 200 people attending each of the 
general excursions into the Gatineau Hills in May 
and September 1902; tickets were 30¢ for adults, i5¢ 
for children, including the cost of the day-long, char- 
tered train trip (LAC OFNC Collection, 13 May 1902). 
Among the participants of the 6 September 1902 ex- 
cursion was Normal School Vice-Principal Sinclair 
“who was accompanied by about 100 Normal School 
students” (LAC OFNC Collection, 6 September 1902). 
One might cynically question if all 100 would have 
been willing participants in their principal’s venture. 

By the early 1900s, however, the Club was not 
nearly so involved in original field-oriented research 
in the Ottawa Valley as it had been one to two decades 
before. There seemed to be a sense that “we pretty 
much know it all now” as research reported in the 
pages of The Ottawa Naturalist grew increasingly more 
national in scope. A greater sense of conservation con- 
cern was developing, however, with the first formal 
conservation action being initiated when the Council 
approved a motion by the Ornithology Committee for 
an OFNC petition calling upon the federal government 
to promote the preservation of shorebird-breeding wet- 
land habitat in the newly established prairie provinces 
(LAC OFNC Collection, 7 April 1908). It was a tenta- 
tive step (an article in that month’s issue of The 
Ottawa Naturalist also provided a prescription for a 
“better” Timber Wolf poison [Anonymous 1908}]). 
Nonetheless, it initiated a long history of conserva- 
tion action and achievement for which the Club has 
been widely honoured and of which the Club can be 
justifiably proud. 

Since its founding, the OFNC had elected a Librarian 
and maintained a natural history library. There were 
regular reports in Council minutes of titles of the in- 
coming scientific literature received in exchange for 
The Ottawa Naturalist or as a professional courtesy 
to individual OFNC members. By 1904, however, it 
was becoming a problem due to the logistical challenge 
of housing and caring for the collection in donated 
space (LAC OFNC Collection, 15 March 1904). In 1906 
the library consisted of “some 350 bound volumes 
occupying about 70 feet of shelf space” as well as 
many unbound volumes (LAC OFNC Collection, 26 
February 1906). The collection seemed to be little 


BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 13 


FEBRUARY, 1901. J 


VOL, XIV, No. 11 


THE 
{| OTTAWA 
NATURALIST. 


Published by the Ottawa Field-Naturalists’ Club. 


CONTENTS. 


x. Annual Address of the President of the Ottawa Ficld- 
Naturalists’ Club selete BHO eipiate 

2. Powers of Adaptation in Fishes, by Professor Edward E. 
Prince 


m 3. Notes of the Acadian Owl (Nyctala Acadica) in captivity, 
by F, Norman Beattie eles steric 4 cows 


(ISSUED JANUARY 23, 


er anne | 


1901.) 


OTTAWA, CANADA, 
Orraws PRintinG Company (Ltwitep). 
3% 5 Moscnrovi Sr. 


FiGuRE 9. The Queen Victoria Memorial issue of The Ottawa 
Naturalist. 


used, however, as the members with significant sci- 
entific questions typically had access to the growing 
institutional libraries at the Museum or “The Farm” 
(as the research centre at the Central Experimental 
Farm was — and still is — known). Arrangements were 
made in 1909 to move the library into the Ottawa Public 
Library where it would be maintained as a stand-alone 
collection (LAC OFNC Collection, 22 June 1909). Pre- 
dictably, the Ottawa Public Library eventually tired of 
the cost and effort of such an arrangement. Faced with 
no other realistic options for its maintenance, in 1917 
the OFNC donated the collection to the OPL, asking 
only that such material be labelled as a Club gift (LAC 
OFNC Collection, 8 March 1917).° Curiously, the posi- 
tion of OFNC Librarian lasted longer than the library, 
being eliminated only in a constitutional change two 
years later (LAC OFNC Collection, 15 January 1919). 

James Fletcher retired from the Council in the spring 
of 1905, after having filled almost every conceivable 
position in the Club and working for it in so many 
other ways throughout its first quarter century. The 
event passed with remarkably little fanfare. Such a 
low-key exit for such a pivotal figure was likely Flet- 
cher’s own doing, however, as indicated by the very 


14 THE CANADIAN FIELD-NATURALIST 


different reaction to his sudden and unexpected death 
on 8 November 1908. 

Although moving along well with an active publi- 
cation, excursions, and environmental education pro- 
gram, the Club was clearly devastated by the loss of 
Fletcher on both an organizational and, for many key 
Club people, a personal level. A number of Club 
founders and luminaries had died earlier, such as 
OFNC 1892-1895 President George Dawson (1849- 
1901), but no one personified the enthusiasm, idealism, 
and commitment of the Club as did James Fletcher. A 
memorial number of The Ottawa Naturalist was pro- 
duced in January 1909, reproducing the many insight- 
ful and heart-felt tributes delivered by professional 
and personal colleagues alike at a special memorial 
event held on | December 1908. The presentations of 
Club co-founders R. B. Whyte and Will Harrington 
were remarkably emotional presentations under such 
stiff, proper Edwardian circumstances. Will Harring- 
ton’s tribute in particular (Harrington 1909) is quite 
capable of tugging heart strings for contemporary 
readers a century removed from that time. With the 
exception of an unfortunately self-promoting speech 
by John Macoun, all contributors that night were clear- 
ly bursting to express their appreciation for the man, 
for his contribution to the Club and for his contribu- 
tion to his chosen country. 

Individual Club members were encouraged to con- 
tribute to a Fletcher Memorial Fund for the construc- 
tion of a memorial drinking fountain to be placed 
near Fletcher’s former work site at The Farm. It was 
quickly oversubscribed, having accumulated over 
$1800 by early 1910 when construction of the memo- 
rial began (LAC OFNC Collection, 15 March 1910). 
A bronze likeness of Fletcher was created by famous 
sculptor (and athletic community associate?) R. Tait 
McKenzie and affixed to the fountain. The fountain, 
with a descriptive plaque attached, was installed in 
1911. It remains along the south side of the NCC 
Scenic Drive in the Central Experiment Farm imme- 
diately east of the Canada Agriculture Museum. 

A memorial painting of James Fletcher was also 
commissioned with surplus funds from the Memorial 
Fund and hung in the Ottawa Public Library? early the 
following spring (LAC OFNC Collection, 12 March 
1912). It was officially unveiled by the federal Minis- 
ter of Agriculture, Sydney Fisher, who “specially dwelt 
on [Fletcher’s] loveable qualities which had endeared 
him so much to all who came in contact with him” 
(LAC OFNC Collection, 19 March 1912). 

It is likely no coincidence that a malaise seemed to 
settle over Club affairs in the following years. Few meet- 
ings of Council or Club excursions were conducted in 
the 1909-1911 period and discussions were held about 
reducing the workload associated with The Ottawa 
Naturalist, including publishing the journal only quar- 
terly.!° The Club operated at a deficit for the first time 
in 1910/1911 (LAC OFNC Collection, 21 March 1911). 
In late 1911 Editor James Macoun reported that the 


Vol. 118 


Club could no longer afford to publish monthly (LAC 
OFNC Collection, 18 December 1911). 

There were calls for — and a dire need of — new 
blood in the organization and President Alexander 
McNeill suggested establishment of “ ... a junior 
branch for the boys. He referred to the success of the 
boy scouts but objected to the military spirit associated 
with it” (LAC OFNC Collection, 12 March 1912). 
Ironic words less than two years before the outbreak 
of World War I. Nothing came of this for many years, 
however, until the founding of the Macoun Field Club 
(see “Post-war Boom (late 1940s-1965)”, below). 

The Ottawa Naturalist seemed similarly to be some- 
what unfocussed, publishing a relatively large number 
of paleontological papers and natural environment 
investigations and reports from afar but with the few 
local contributions increasingly confined to popular 
topics. A review article on the horrors of Poison-ivy 
allergic reactions provides an example, reminding 
any potentially afflicted reader that “ ... nothing is 
better than the old-fashioned lead and opium lotion 
...” to relieve the itching (Macnamara 1912). 

The notable exception to this was the effort to devel- 
op an active environmental protection and bird con- 
servation program, led by Gordon Hewitt (1885-1920). 
Hewitt was Fletcher’s replacement as Dominion Ento- 
mologist (in 1909) and like him, was a dynamic, per- 
sonable English immigrant (Criddle 1920). During 
his few years in Canada he played an important role 
in North American natural sciences and conservation 
matters, including being the lead Canadian official in 
the development of the 1916 Migratory Birds Con- 
vention, before dying at a tragically young age in the 
post-war influenza epidemic (Foster 1978). Likely be- 
ing assisted by his socially prominent position as the 
husband of Prime Minister Bordon’s niece, he suc- 
cessfully lobbied both the Ottawa Improvement Com- 
mission (forerunner of the National Capital Commis- 
sion) and the Boy Scouts of Canada to become in- 
volved in enhancement of migratory bird habitat (LAC 
OFNC Collection, 17 February 1914). His crushing 
load of professional responsibilities, however, limited 
the amount of time he was able to put into re-ener- 
gizing the Club, even during his tenure as President 
in the difficult war-time period of 1918-1919. 

As with so many cultural, social, and technical insti- 
tutions in Canada, the OFNC was devastated by events 
surrounding Canadian involvement in World War I. 
On top of their post-Fletcher era struggles, the burden 
of war-time logistics, costs, and priorities came close to 
destroying the OFNC. The Club ran deficits through 
the war years and for the first time in Club history, the 
steady rate of membership growth virtually stopped 
(Figure 7). A new focus and a new source of energy 
were desperately needed. They came in the form of 
another recent arrival in Ottawa, this one being a tall, 
quiet, bearded architect turned ornithologist from 
southwestern Ontario who would turn the OFNC into 
a national institution. 


2004 


A National Role (1918-early 1940s) 

Percy Taverner (1875-1947) didn’t think much of 
Ottawa or Ottawa naturalists when he arrived in 1911 
as the National Museum’s first ornithologist (Cranmer- 
Byng 1996). Shortly after arriving he complained to a 
Detroit friend, “we have a club here, the Ottawa Nat- 
uralists, who are much on a par with the Detroit bunch. 
They have nice picnics every week but they are no 
place for you and me. The worst of it is that they 
have all kinds of direction with good men. Fletcher, 
Macoun, Gibson and the whole Geological Survey and 
that of the Experimental Farm, but they haven’t evolved 
a single naturalist in their twenty-five years of existence. 
The only thing they have got is a publication that has a 
government grant and appears regularly and in which 
we can get publication any time. If not for that, the 
real students here would have let the whole organiza- 
tion die a natural death long ago”. (LAC Taverner Col- 
lection, 29 April 1912). 

Taverner’s criticism was brutal and a bit unfair — 
but not by much. And to both his eternal credit and 
the benefit of Canadian natural sciences, he set about 
working with other like-minded rebels to shake the old 
outfit up and to make it — or at least, its publication — 
better serve the needs and opportunities of the contem- 
porary naturalist and public communities. Fletcher, 
Martin, Henry B. Small, Whyte, and Harrington would 
have been proud. 

Like Fletcher before him, Taverner maintained an 
extensive network of correspondents across Canada 
and the United States. Although allied on the OFNC 
Council with the likes of botanist James Macoun, long- 
time editor of The Ottawa Naturalist, entomologist 
Arthur Gibson (1875-1959), and herpetologist Clyde 
Patch (1887-1952), Taverner was inspired by his field- 
naturalist colleagues across the country to broaden 
the reach of the Club’s publication. Early in 1918 he 
argued to the Council that “... it does not seem pos- 
sible to support a worthy publication in a purely local 
field. Any such endeavours to be successful must en- 
large its field. I therefore suggest that the pure local 
character of the periodical be removed by a change 
of name and propose The Field Naturalist” (LAC 
OFNC Collection, 26 March 1918). 

Taverner, Macoun, and associates made the skepti- 
cal Council an offer they could not refuse. They prom- 
ised it would mean no substantive change for local 
naturalists and would still accommodate more popu- 
lar general public issues. Most importantly, they per- 
sonally guaranteed to underwrite a huge expansion in 
membership in order to establish a secure, long-term 
financial base for the program. James Macoun, for ex- 
ample, immediately pledged to underwrite 100 new 
memberships; Taverner, Patch and anthropologist F. 
W. Waugh each took on 25 and archaeologist W. J. 
Wintemberg accepted responsibility for a further 15 
(LAC OFNC Collection, 1 May 1918; 17 March 1919). 
How could the Council refuse such an offer? 


BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 15 


Vor. XXXII, No. 1. APRIL, 1916. 


aS 


| 


second ebzss malice. 


FiGure 10. An issue of the last volume of The Ottawa Nat- 
uralist in 1918. 


Taverner described the manoeuver to his mentor J. 
H. Fleming in August 1918 as something of a coup, 
“ .. a few of us got in control and overrode the con- 
servatives ... ” (Cranmer-Byng 1986). He went on in 
the Fleming letter to say, “we hope to make it the sci- 
entific and nature study periodical of Canada” (LAC 
Taverner Collection, 10 August 1918). And indeed, 
by the following spring the mailing list had virtually 
doubled to over 540 names (LAC OFNC Collection, 
17 March 1919). Taverner reported back to a skepti- 
cal Fleming in Toronto that “you are mistaken in 
believing that the change in name will not help The 
Ottawa Naturalist. It has helped already. The change 
in form can only be supported by increased subscrip- 
tions and we have to rely mostly on Canadians for this. 
The jealousy of anything labeled Ottawa throughout 
Canada is surprising” (LAC Taverner Collection, 10 
April 1919). 

The Canadian Field-Naturalist (CEN) was officially 
established by a motion from James Macoun (sec- 
onded by Taverner) at the OFNC Annual Meeting in 
March 1919. A change had already been made in April 
1918 with a larger format and better paper (Figure 10), 


16 THE CANADIAN FIELD-NATURALIST 


graced by the attractive, if pastoral, cover illustration 
provided by Geological Survey of Canada artist C. E. 
Johnson. The new journal name was first used on the 
April 1919 issue. This cover illustration remained on 
each regular issue of the CFN until volume 59 (1945) 
when, without further explanation, it was removed in 
favour of the Table of Contents (LAC OFNC Collec- 
tion, 3 March 1945). 

The care and maintenance of the CFN became the 
primary issue and function of the Club. The under- 
writing of new subscriptions by the small group of 
‘friends’ of the journal had a dramatic effect on mem- 
bership numbers, the new levels of which persisted 
through the 1920s (Figure 6). This laid a strong, nation- 
al foundation for the publication without which it seems 
unlikely the publication (and the OFNC) would have 
survived the dark days of the 1930s. 

Another major publication initiative in that period 
was the first OFNC Special Publication, occasioned 
by the death of John Macoun. Upon the urging of his 
son, horticulturist and OFNC 1903-1905 President 
William T. Macoun (1869-1933), the Council agreed 
to publish the elder Macoun’s autobiography (LAC 
OFNC Collection, 9 April 1921). Curiously, not since 
the Ottawa Natural History Society produced its occa- 
sional Transactions more than 50 years earlier had 
such a stand-alone publication been presented by the 
Ottawa naturalist community. 

While a Memorial Fund was established to cover 
the $2500 costs of publication of the Macoun autobi- 
ography as well as a memorial portrait (again as with 
James Fletcher’s portrait, created by Franklin Brown- 
ell), it apparently fell to William Macoun to make 
most arrangements (LAC OFNC Collection, 9 April 
1921). This he did enthusiastically and efficiently, lead- 
ing to an initial run of 2000 copies of the autobiog- 
raphy (Macoun 1922). Indeed, after the book was pub- 
lished, the 1922 OFNC Annual Report was deliberately 
altered to read that “the Club had co-operated with Mr. 
W. T. Macoun in the publication of the Autobiography 
of Professor John Macoun” (LAC OFNC Collection, 
5 December 1922), suggesting that the Club may not 
have played much of a role in its development."! 

The Macoun portrait was formally presented to Na- 
tional Museum of Canada Director William McInnis 
during the 1921 OFNC Annual Meeting which was 
held in the Victoria Memorial Museum, Macoun’s last 
work place in Ottawa. It was accompanied by various 
speeches praising his extraordinary contributions to 
Canadian natural science (LAC OFNC Collection, 20 
December 1921). 

Other occasional publications appeared as Special 
Issues of the CFN through the 1920s. One was a treat- 
ment of the birds of Saskatchewan which included the 
first colour illustration to appear in the journal (Mitch- 
ell 1924) and another was a long essay on the natural 
resource potential of northern Canada (Kindle 1928), 


Vol. 118 


copiously illustrated by high quality black-and-white 
and half-tone plates. The latter, by paleontologist and 
OFNC 1927-1928 President E. M. Kindle (1869-1940), 
who had recently won a $1000 prize in a Canada-wide 
competition established by Sir William Price for the 
best article on this subject. This pattern of occasional 
Special Issues for larger, particularly significant or 
appropriate subjects has continued to the present day, 
recent subjects including the Taverner biography 
(Cranmer-Byng 1996) and a review of the orchid spe- 
cies of the Ottawa District (Reddoch and Reddoch 
1997). Earlier long articles, like Fletcher’s Flora 
Ottawaensis (Boivin and Cody 1955) or Hoyes Lloyd’s 
review of the birds of Ottawa (Lloyd 1923; 1924), 
were often serialized over a number of issues. 

Another special publication was called for in the 
1920s. It was recommended to the Council by Geo- 
logical Survey of Canada geologist J. B. Mawdsley 
that the Club produce “ ... a local scientific guide book 
of the region” (LAC OFNC Collection, 20 November 
1929). The OFNC was indeed instrumental in seeing 
such a naturalist’s guide produced, but that did not 
happen until nearly 60 years later (Brunton 1988). 

Local programming became little more than occa- 
sional field outings and lectures during the late teens 
and 1920s, although efforts were made to get the Boy 
Scouts organization more involved during the early 
1920s (LAC OFNC Collection, 26 March 1921). 
Nonetheless, Taverner’s earlier unflattering portrayal 
of the Club’s activity level was still more or less on 
target a decade later. This apparent lack of new energy 
and focus in the post-war years and early days of the 
Roaring Twenties was underscored by the deaths of a 
number of prominent OFNC pioneers and activists, 
men who had emphasized the need for and impor- 
tance of local field activity. These included founders 
R. B. Whyte (1919) and Will Harrington (1918), as 
well as Gordon Hewitt (1920), James Macoun (1920), 
and John Macoun (1920). 

National conservation issues, typically relating to 
the protection of particular animal populations, con- 
stituted the major non-publications subject of discus- 
sion of the Council in the 1920s and 1930s. On a num- 
ber of occasions the Council passed motions for the 
Club to petition the federal government regarding such 
issues as improved national museum support or the 
protection of wildlife. This may have been awkward at 
times, since many of the people receiving and dealing 
with such pleas or critiques were Club members or 
professional associates of Club members. Senior gov- 
ernment eyebrows may have been raised, for instance, 
when Editor Harrison Lewis (1893-1974) published a 
(prescient!) editorial in the CFN that was highly criti- 
cal of apparent governmental support for increased 
commercialization of Canadian national parks (Lewis 
1922). It has never been, after all, particularly judi- 
cious for active civil servants to publicly question 


2004 


stated government policies or intentions. “Awkward”, 
however, does not adequately describe the “Buffalo 
Crisis” of 1925. 

The trouble arose when the Council directed that a 
letter be sent to the Department of the Interior protest- 
ing the proposed movement of Plains Bison of ques- 
tionable health into the disease-free range of the last 
known herd of Wood Bison (LAC OFNC Collection, 
28 February 1925). The protest letter was signed by 
OFNC President Hoyes Lloyd (1888-1978) and accom- 
panied by a comparably critical Letter to the Editor 
which had recently been published in The Canadian 
Field-Naturalist (Harper 1925). Lloyd and the CFN 
Editor Harrison Lewis, however, were both employees 
of the Department of the Interior and their public crit- 
icism of their professional superiors was clearly not 
appreciated. Instructed to choose between continued 
federal government employment and their OFNC 
positions, both Lloyd and Lewis had little choice but 
to resign their Club positions immediately — to the 
apparent surprise and regret of the Council (LAC 
OFNC Collection, 11 April 1925). Lloyd became the 
first and only OFNC President forced to resign in mid- 
term, although he served the Club in many ways for 
decades thereafter and was awarded an Honorary Mem- 
bership in 1965 in recognition of his contributions 
(Munro 1979). Curiously, the taking of this laudable, 
if perhaps somewhat naive position of principle, was 
publicly unreported until 2002. It is not even hinted 
at in The Canadian Field-Naturalist obituaries of either 
gentleman (Solman 1974; Munro 1979). Lewis, Lloyd, 
Harper and associates have been proven correct in their 
concerns regarding herd contamination, as the issue of 
appropriate Wood Bison population management in 
Wood Buffalo National Park is once again being hot- 
ly debated (Fuller 2002). 

On a more positive note, the 50“ anniversary of 
the OFNC was celebrated with an anniversary dinner 
on 19 March 1929 (erroneously believed to be the 
founding date — see “Founding Meeting”, above), in 
which two of the few surviving founders (H. B. Small 
Jr. and Roger Davy) were the guests of honour. Other 
founders (participants in the 25 March 1879 found- 
ing meeting) still alive in March 1929 were L. H. 
Chrysler, W. Chesterton, and P. B. Symes (who died 
later that year).!* The Club was doing reasonably well 
in early 1929, however, with a substantial surplus and 
a stable, adequate number of members to support its 
programs. If local activities were not undertaken at 
anything like the keen levels of years and decades 
before, the publication program seemed solidly and 
comfortably established and seemed to have achieved 
the national scope that Taverner, James Macoun, Lewis, 
and their associates had worked so hard a decade ear- 
lier to achieve. 

The respite from difficult times was short-lived, 
however, as the Great Depression of the 1930s pre- 


BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 17 


sented another severe test of the Club’s staying power 
and its officers’ mettle. Stripped down to basic func- 
tions — publishing the CFN and little else — the Club 
was enduring especially tough times, ushered in by 
dramatically falling membership in 1930 and a sub- 
stantial budget deficit. There was a $728 swing in net 
revenues from the surplus of the previous year (LAC 
OFNC Collection, 11 April 1930). Membership would 
not again exceed the 1929 total of 549 until 1955 ( 
6). 

By the December 1932 Annual Meeting the OFNC 
was operating at a 20% deficit — and this only after 
drawing upon the Club’s small Reserve Fund and 
securing a 12% reduction in charges from the CFN 
printer. The good news, though, was that excursion 
attendance was way up, with four spring field trips 
each averaging over 100 attendees. OFNC 1931-1933 
President Charles Sternberg (1885-1981) concluded 
cheerfully “we feel our organization has stood up as 
well as others in the stormy times and we are, as a 
Club, now looking forward and ready to face brighter 
days as they dawn upon us” (LAC OFNC Collection, 
6 December 1932). 

It was not to be, of course. The Club officers were 
creative and flexible in finding ways of keeping things 
going despite the depletion of the Reserve Funds and 
low membership numbers. They agreed, for instance, 
to carry the 1934 membership of anyone who had been 
with the Club for five years but could not afford the 
$2.00 membership/subscription fee (LAC OFNC Col- 
lection, 9 January 1934). Local excursions were em- 
phasized again (e.g., 10 were held in 1935) and while 
these did not appear to generate much in the way of 
new biodiversity information, they did offer a benefit 
and encouragement to local members. A re-emphasis 
on conservation matters was also expressed, especially 
by those working in National Parks and in federal 
migratory bird conservation offices, but no particular 
hands-on local initiatives were identified. The Council 
even decided to defer joining the newly established 
Federation of Ontario Naturalists which was dedicated 
to the objective of protecting natural areas and “native 
wildlife” (biodiversity) in Ontario (LAC OFNC Col- 
lection, 7 May 1936). 

Despite the difficult times and reduced level of local 
activity, the 1936 Annual Business Meeting drew over 
100 members, likely due to the attendance of Archie 
Belaney, a.k.a. Grey Owl (Dickson 1973) “... Canada’s 
famous Indian naturalist and conservationist who 
spoke briefly on the need for immediate conservation 
action in Canada” (LAC OFNC Collection, 1 Decem- 
ber 1936). 

What sustained the OFNC through all the ups and 
downs of this difficult time was the special relation- 
ship of the Club with the community of biologists 
and scientists employed by the federal government, 
especially with those at The Museum (Geological 


18 THE CANADIAN FIELD-NATURALIST 


Survey of Canada Museum, then National Museum 
of Canada, now Canadian Museum of Nature) and 
The Farm (Central Experimental Farm, now various 
research initiatives within Agriculture Canada). Hall 
(1986) notes that over 30 employees of The Farm — 
the most notable being James Fletcher — served as 
officers, editors, and/or Honorary Members of the 
OFNC (Appendix 1). Twelve of those individuals had 
been OFNC presidents. A comparable or possibly even 
stronger commitment could be documented from The 
Museum, with Department of the Interior (Canadian 
Wildlife Service (CWS) and Parks Canada) personnel 
making a similarly important contribution to the opera- 
tion and support of the OFNC. These people saw the 
Club in general and the CFN in particular as an im- 
portant contribution to Canadian and North American 
natural sciences and to the conservation of the natur- 
al environment. They saw it (as many in such positions 
still do) as virtually their duty to participate. In an 
otherwise mundane 1937 debate on the question of 
possible reduction in the honorarium given to the CFN 
editor (from the princely of $90 established in 1928, 
to $50) as yet another cost-cutting measure, Harrison 
Lewis stated “... we are a scientific society and any 
one of us would carry on the work without salary for 
the good of the Club”, to which Percy Taverner 
added ” ... the Editor works for the good of his 
science and not for wages ...”. The motion for a 
reduction in the editor’s honorarium passed, by the 
way (LAC OFNC Collection, 10 April 1937), and was 
not increased again until 1947 (LAC OFNC Collec- 
tion, 25 January 1947). 

There remained a small group of Club officers 
throughout the 1930s and early 1940s who argued for 
more emphasis on a strong local program of lectures, 
excursions, and public education. This group was led 
by irrepressible biology teacher and priest F. E, Banim 
(1902-1979), statistician W. H. Lanceley (1893-1958), 
and technical editor Pauline Snure. These Club offi- 
cials, also supported by botanist and OFNC 1935-1937 
President Herb Groh (1883-1971), argued for the re- 
establishment of monthly lectures and a return to old 
time soirées as means of generating renewed interest 
and membership in the Club. They were successful in 
the establishment of a Flora and Fauna Committee to 
co-ordinate a renewal in the investigation and docu- 
mentation of local biodiversity (LAC OFNC Collec- 
tion, 28 October 1936). This latter committee ultimate- 
ly had few results, although J. R. Dymond, selected as 
leader of the fish working group, presumably conduct- 
ed at least a part the research which was the basis for 
his later The Fishes of the Ottawa region (Dymond 1939) 
in response to this initiative. 

Snure was particularly forceful in the mid-1940s 
in taking on long-established, senior Council members 
like Hoyes Lloyd and, to a lesser extent, Harrison 
Lewis and Percy Taverner, and pushing successfully 
for such things as the establishment of an Excursions 
and Lectures Committee to co-ordinate services for 


Vol. 118 


local members (LAC OFNC Collection, 22 January 
1944). The latter three — now old guard members — 
were the most vocal on Council in arguing for contin- 
ued priority being placed on the important “national 
publication society” elements they had worked so 
hard to sustain through the difficult 1930s and into 
the early 1940s. Although their efforts only slowly 
bore fruit, Banim, Lanceley, and Snure (who served 
consecutively as OFNC President between 1944 and 
1950)!°, were able to shepherd a measure of renewal 
of local naturalist activities in this period. 

The at-times conflicting perspectives of local natur- 
alist club and national scientific publication society 
simmered along, unresolved as national and world 
events (the latter years of the Great Depression, the 
commencement of World War II) effectively limited 
the ability of the Club to generate substantial new 
membership or financial resources. Despite the war- 
time limitation on materials and human resources, 
some Club local field investigations were initiated 
and the CFN continued to appear, albeit with many 
issues in 1941 and 1942 delayed by the printer’s war 
work priorities. Not oblivious to patriotic opportuni- 
ties themselves, in 1942 the Council responded to a 
request for the exchange of scientific literature from 
the Lenin Library in Moscow by offering a free set of 
The CFN from 1935 “until the close of hostilities ... 
as a mark of our good will” (LAC OFNC Collection, 
5 October 1942). For OFNC functions and programs, 
the World War II period was essentially an extension of 
the publications and membership subsistence efforts 
of the 1930s. Only as the war in Europe was winding 
down did the first signs of renewal and new potentials 
— such as increased membership — begin to appear 
(LAC OFNC Collection, 5 December 1944). 


Post-war Boom (late 1940s-1965) 

One of the first signs of the rebirth of local area 
activity within the OFNC came with publication of 
review articles on the birds and mammals of the 
Ottawa District in the latter years of World War II 
(Lloyd 1944 and Rand 1945, respectively), at least in 
part at the urging of local naturalist advocates like 
Banim, Snure and Lanceley. Although based largely on 
historic Museum data and/or the authors’ observations, 
these review papers gave local naturalists the first com- 
prehensive treatments of popular Ottawa District flora 
or fauna since Fletcher’s 19" century vascular plant 
list (Fletcher 1888) and Lloyd’s bird list from the 
1920s (Lloyd 1923; 1924). 

Almost coincident with, but much more important 
than this, was the tremendous post-war growth in the 
professional civil service in response to the needs of 
an expanding, prosperous post-war national economy. 
A new wave of young, energetic, natural scientists 
from all across Canada was arriving in Ottawa in the 
late 1940s and early 1950s, many soon joining and be- 
coming involved with the programs and workings of 
the OFNC. Following the dark days of the Depression 


2004 


and then the war, many of these individuals were keen 
to contribute both to the improvement of their recover- 
ing local and national communities and to the explor- 
ation and protection of Canadian natural biodiversity. 
In November 1946, for example, future Honorary 
members and important Club contributors Clarence 
Frankton (1906-2000) (Brunton 2003) and Jack Gillett 
joined the Club, followed the next month by long-time 
Business Manager of the CFN and future Honorary 
Member Bill Cody. Indeed, almost 150 new members 
joined the OFNC in 1948 alone (Snure 1978). Newly 
arrived natural science specialists were pressed by 
their OFNC veteran associates to join the Club; in- 
volvement in Club programs was almost an expected 
part of the job in those years (C. Frankton, personal 
communication). 

This new wave of naturalists demonstrated some- 
thing not seen since the early years of the Club’s his- 
tory — a keen enthusiasm for field exploration and 
discovery, and a desire to share that new-found know- 
ledge with the naturalist community and the general 
public. Accordingly, the late 1940s and early 1950s 
were productive years for the documentation of Ottawa 
area biodiversity, as indicated by the richness of the 
specimen records from that time in the collections of 
The Museum and The Farm (personal observation). 

Another major initiative of this period was the 
formation of the Macoun Field Club (MFC), a junior 
naturalists club sponsored jointly by the OFNC and 
the National Museum of Canada (Snure 1978; Baldwin 
1978). Such an organization had been talked about 
since before the First World War, but nothing had 
come of it. Due to dedicated work by a large number of 
Club and Museum people, however, it was success- 
fully established in 1948. The first MFC Committee 
Chairman was OFNC 1954-1955 President W. K. W. 
(Bill) Baldwin (1910-1979) (Figure 11) who set the 
initial tone and procedures which have stood up ever 
since (Soper and Bousfield 1982; Francis Cook, per- 
sonal communication). The MFC has supported and 
encouraged the investigation and documentation of nat- 
ural biodiversity by elementary and secondary school 
children in the Ottawa area through a remarkably rich 
program of lectures, workshops and most importantly, 
field studies, for over 50 years (Lee 1998). The work of 
Bill, Herb Groh (Figure 12) and David Maddox was 
particularly critical to this success in the formative years 
of the MFC. Many OFNC club members contributed 
time and even, in the case of Mary Stuart, access to their 
property for field trips (Figure 13). 

The OFNC also undertook to generate greater pub- 
lic awareness of natural environment matters and to 
develop an additional stream of funding for the Club 
by arranging local sponsorship of Audubon Screen 
Tour presentations in Ottawa. These were profession- 
ally produced and well-attended illustrated lectures 
by speakers of the calibre of Roger T. Peterson and 
George M. Sutton (personal observation). They pro- 
vided an important source of general public education 


BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 19 


on a wide range of natural environment topics in the 
days before specialty television channels or, indeed, 
television of any kind was available for most Ottawa 
households. It’s not entirely clear who was the Club’s 
lead on this program; Humphreys (1979a) suggests it 
resulted from Oliver Hewitt’s circulation of an Audu- 
bon Screen Tour brochure in December 1947. The rec- 
ords of the minutes books, however, state that former 
Treasurer and future Honorary Member Ibra Connors 
(1894-1989) brought the issue before the Council for 
approval later that winter (LAC OFNC Collection, 21 
February 1948). Regardless, until it was cancelled in 
1959, the Audubon Screen Tours program was enjoyed 
by many thousands of Ottawans and generated con- 
siderable revenue for the Club (Humphreys 197%a). 

A significant amount of Club revenue also resulted 
from the clearance of a mass of The CFN back issue 
orders which had built up during the 1941-1955 edi- 
torship of Harold Senn (1912-1997). Bill Cody and 
Clarrie Frankton spent weeks ferreting out and follow- 
ing up on years-old orders for The CFN back issues 
(C. Frankton, personal communications). The resulting 
revenue formed the seed funding which has grown into 
what now constitute very substantial OFNC reserve 
funds. 

In the late 1940s and for the first time in OFNC 
history, the Club had its own field station. In 1949 A. 
E. (Fred) Bourguignon (1893-1968) arranged for a 
10-year lease of a property at Beatty Point along the 
Ottawa River off what is now Grandview Road in 
Nepean. For a cost of $300 (assisted by a substantial 
donation of building materials), various Club members 
(most particularly, Fred Bourguignon) worked to 
construct a small wooden building which would serve 
as a base for OFNC field studies in the area. Beatty 
Point Lodge, as it was called, was officially opened 
on 24 May 1949 and served as a field investigation 
centre for almost 10 years. Encroaching urban growth, 
expanding road networks, and the proliferation of per- 
sonal automobiles eventually made a single base of 
operations less useful for contemporary field explor- 
ations and the building was sold in the late 1950s 
(Humphreys 1979b). 

Some members in the early 1950s wished to delve 
more deeply into particularly popular natural history 
subject areas in the Ottawa District. From this interest 
arose informal study groups such as the Fern Group, 
the Bog Group, the Trail Group, the Bird Group, etc. 
(Dill 1979; 1982), and later, the Native Orchid Loca- 
tion Survey. Consistent with the science-based approach 
of OFNC field investigation throughout its history, 
these study group participants did more than just ex- 
plore interesting natural landscapes and share identi- 
fication information amongst themselves. They docu- 
mented significant findings in writing and/or with spec- 
imens. The records of the Fern Group, for example, 
were heavily relied upon for the production of a 
treatment of the ferns of the Ottawa District (Cody 
1956). The Native Orchid Location Survey was estab- 


20 THE CANADIAN FIELD-NATURALIST Vol. 118 


Ficure 11. Bill Baldwin, botanist at the National Museum of Canada and the first chair of the Macoun Field Club, at the meet- 
ing room provided by co-sponsor National Museum of Canada in the basement of the Victoria Memorial Museum 
Building together with early members Cynthia Millman and Nancy Fergusson, November 1950 (Macoun Club files, 
courtesy of Rob Lee). 


FiGurE 12. Herb Groh, then just retired weed specialist at the Canada Department of Agriculture and a Past President of The 
Ottawa Field-Naturalists’ Club, with Macoun Field Club member Nick Wickenden, November 1950. (Macoun Club 
files, courtesy of Rob Lee). 


2004 


BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB aM 


FiGurE 13. One of many Macoun Field Club trips to OFNC Honorary Member and long-time Macoun volunteer Mary 
Stuart’s property near Packenham: Intermediate and junior groups, March 1969. Photograph courtesy of Rob Lee. 


lished by E. W. Greenwood in 1966 and soon grew in 
scope beyond the Ottawa District, gathering a consid- 
erable body of orchid location data from volunteer 
contributors across Canada until the mid-1970s (Red- 
doch and Reddoch 1997). 

With local natural history activities reaching levels 
not seen since the early days of the century, OFNC 
members felt the need for more timely and topical 
information through a forum which would help to 
“tie the members together” (LAC OFNC Collection, 
28 February 1949). Appropriately enough, that initia- 
tive was approved by the Council at the same meeting 
authorizing the construction of the Beatty Point Lodge, 
underscoring the new-found enthusiasm for local activ- 
ity. The first issue of the Ottawa Field-Naturalists’ 
Club Newsletter was published in April 1949 under 
the editorship of Verna (Ross) McGiffin. It ran until 
1967, providing timely notices of meetings and events 
and offering a forum for the speedy exchange of in- 
formation and opinions on a wide variety of issues of 
interest to the local membership. 

The Club reached its 75" Anniversary in 1954, 
though due to program delays, the occasion was actu- 
ally celebrated in 1955 (Dill 1979). It was much more 
of a party affair than the low-key Founders’ Tea and 
Club dinner held for the 50" anniversary back in 1929. 


The Club was profiled in the print media and a local 
department store provided a display window for an 
exhibit on the Club. The attention paid to the event 
generated the first formal demonstration of historical 
interest in the organization. Herb Groh, whose OFNC 
involvement dated back to the pioneer days of the 
Club and who had worked with both John Macoun 
and James Fletcher (Taschereau 1972), provided an 
anecdotal history of Club Presidents (Groh 1955). 
After all, he’d known almost all of them! Although 
some historical documentation of technical matters 
pertaining to the CFN’s publication history had been 
produced (LaRocque 1931; Boivin and Cody 1955), 
this was the first public documentation of the person- 
alities and events that had highlighted the Club’s 
history.'* 

The CFN produced increasingly larger issues 
through this period, reflecting the greater volume and 
quality of original field work being conducted both 
locally and farther afield in the early to mid-1950s. 
This increased production, however, posed financial 
problems as publication costs increased 400% from 
the mid-1940s to 1953; Club income not quite dou- 
bled in the same period. Since it was felt that mem- 
bership fees ($2.00) were as high as could be sustained, 
costs had to be reduced. Accordingly, after decades 


22 THE CANADIAN FIELD-NATURALIST 


TRAIL, & 
wi ye 


A PUBLICATION CONCERNED WITH 
NATURAL HISTORY AND CONSERVATION 


TRAIL & LANDSCAPE Vol.1.No.1 p. 1-28 Ottawa, March-April, 1967 


FiGure 14. The first issue of Trail & Landscape, March-April 
1967, a journal for regional observations and outings 
and Club events. The cover was designed by the first 
editor, Anne Hanes. 


of debate on the matter, the journal was reduced to 
quarterly publication (from six issues per year) ef- 
fective with volume 67 (1953) (LAC OFNC Collection, 
20 November 1952). Publication production problems 
were seriously exacerbated by lengthy delays in getting 
into press, leading Publications Chairman and OFNC 
1950-1952 President Walton Groves (1906-1970) to 
“express his deep regret at his inability to command 
the support and co-operation” of just retired editor 
Harold Senn (who had served 13 % years) in order to 
pass on the editorship to incoming editor R. A. Ham- 
ilton (LAC OFNC Collection, 4 May 1956). By the 
end of 1956 the last issue of the 1955 volume had still 
not been produced. The Council decided not to pay out 
the editor’s honorarium for that year until the volume 
was completed (LAC OFNC Collection, 13 December 
1956). The crisis was resolved by a flurry of publica- 
tion activity under the efficient new editor, however, 
with seven issues being published between February 
and September 1957 (LAC OFNC Collection, 30 
September 1957)!'> 

The Club continued to struggle financially despite 
the generally rosy economic picture in the country, as 
it also had, ironically, during the “Roaring Twenties”. 


Vol. 118 


In the mid-1950s, for example, Treasurer R. J. Moore 
advised the Council “to go easy, especially keep 
Excursions & Lectures Committee under control” 
(LAC OFNC Collection, 29 February 1956). Walton 
Groves’ assessment was even more brutal, stating that 
“many of the Club’s activities have been crippled by 
the delay in publication” and by the need to publish 
sO many issues in such a short time (LAC OFNC 
Collection, 30 September 1956). Despite the return to 
timely publication of the CFN, at the end of the decade 
Groves still reported that “the financial position of 
the Club continues to be very grave” at (LAC OFNC 
Collection, 1 December 1960). 

Still, the OFNC managed a Special Issue of The 
CEN, Alice Wilson’s Geology of Ottawa (Wilson 
1956). It was likely only possible because Carleton 
University placed a pre-publication order for 1000 
reprints (LAC OFNC Collection, 13 April 1956). 

Before environmental conservation and the protec- 
tion of natural biodiversity became major social and 
cultural phenomena in the Western World in the 1960s, 
the locally-focused elements within the OFNC had 
been gradually increasing the organization’s level of 
involvement in such directions. In the early 1950s, 
for example, Harrison Lewis met with the Federal 
District Commission (FDC) to discuss how the OFNC 
could (and would) provide wildlife protection and 
enhancement consultation during Gatineau Park dev- 
elopment. They also explored the idea of an OFNC 
field station space being provided in an expropriated 
Gatineau Park building (LAC OFNC Collection, 7 
April 1951). An ORNC Gatineau Park Committee was 
formed under the chairmanship of OFNC 1960-1961 
President Winston Mair and made various recom- 
mendations for wildlife habitat enhancement there. 
The Federal District Commission also asked the Club 
to be involved in the establishment of a Brewery Creek 
bird sanctuary in Hull (Gatineau), presumably in res- 
ponse to the high profile that site had received from 
former British High Commissioner Malcolm Mac- 
Donald’s war time reminiscences in Birds of Brewery 
Creek (MacDonald 1947) (LAC OFNC Collection, 
13 March 1952). Nothing came of that and neither 
FDC consultation was mentioned again after 1954 
(LAC OFNC Collection, 15 December 1954). 


Conservation priorities (late 1960s-early 1980s) 

A forerunner of the OFNC Conservation Commit- 
tee, called the Preservation of Natural History Sites 
Committee, was established in 1960 with Bill Baldwin 
as its chair and with locally-active members including 
biology professor Donald A. Smith, CWS biologist 
Vic Solman, geologist D. D. Hogarth and botanist W. 
G. Dore (1912-1996) (LAC OFNC Collection, 15 
December 1960). One of its first ventures — opposing 
the proposed destruction of the vast Mer Bleue peat- 
land in eastern Ottawa (it was to be used as a regional 
landfill!) — was highly successful. It led not only to 


2004 


BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 23 


FIGURE 15. Members of the Traill Group near Pinks Lake 29. May 1965, one of the last outings before disbanding as a 
formal group. From left to right are Winifred Anderson, Ruth Resenel, Alice Frith, Sheila Thomson, Bill Thomson, 
Rowley Frith, Hue McKenzie, Elva MacKenzie, Anne Hanes and unknown. Photograph by Charlotte Dill. Trail & 
Landscape 16(2): 95 [1982]. 


protection of that magnificent natural area, but also 
to a series of ecological and biodiversity investigations 
of the wetland (Baldwin and Mosquin 1969). Donald 
Smith played a pivotal role in generating the neces- 
sary awareness amongst both the naturalist commu- 
nity and National Capital Commission (NCC) officials 
of the significance of the bog (Dorais et al. 1974). 
The protected area — now some 3 500 ha in size — is 
managed by the NCC for the benefit of its ecological 
functions and has been declared an internationally 
significant wetland under the Ramsar Convention on 
Wetlands (National Capital Commission 1996). 

In this period local Club members and the general 
public benefitted from two birding columns in Ottawa 
newspapers, one in The Ottawa Citizen by Wilfred 
Bell and the other in The Ottawa Journal by the ap- 
propriately named John Bird. The latter weekly column, 
“Bird’s-eye View”, was particularly beautifully written, 
accurate and insightful. It emphasized the growing 
level of birding activity in the Ottawa Valley and 
offered a timely (unofficial) bulletin board of OFNC 
and conservation events (McNicholl 1994). 

Certainly the most important single occurrence for 
the local Club element in this period was the estab- 
lishment of Trail & Landscape (T&L) (Figure 14). 
As expressed by the then CFN editor Theodore Mos- 


quin, it was to be “a newsletter as well as contain[ing] 
articles of wide appeal in the conservation field” (LAC 
OFNC Collection, 5 January 1967). The existing OFNC 
Newsletter wasn’t offering sufficiently comprehensive 
local coverage, nor could it satisfactorily accommodate 
the documentation of local conservation-oriented nat- 
ural environment investigations which were increas- 
ingly being undertaken by Club members. T&L was 
an immediate and astonishing hit, not only satisfying 
a need of active local Club members but generating a 
huge, unprecedented increase in membership (Figure 
6). Anne Hanes (1925-1981) was appointed as the first 
editor and exercised these duties superbly through 13 
volumes (1967 to 1979), always encouraging environ- 
mental conservation themes wherever possible (Green- 
wood 1980). 

Two subsequent editors, Joyce Reddoch (from 1980 
to 1989) and Fenja Brodo (from 1991 to 1992 and 1993 
to 2001) expertly supervised and directed the develop- 
ment of long runs of the publication. Reddoch’s term 
was distinguished by the production of a number of 
large review articles which she solicited for both their 
intrinsic natural history value and their importance as 
reliable references for various conservation applications 
thereafter. A number of these, such as the reviews of 
Ottawa District amphibian and reptile species (Cook 


24 THE CANADIAN FIELD-NATURALIST 


1981, a revision of an earlier series in 1967 solicited by 
first editor Anne Hanes) and butterfly species (Layberry 
et al. 1982), continue to be locally referenced, as do 
all of the natural areas documentation published in 
T&L. The Club and its members have also been well 
served by other editors of Trail & Landscape, Elizabeth 
Morton (1990), Bill Gummer (from 1992 to 1993) and 
Karen McLachlan/Hamilton (from 2001 to the present). 

Trail & Landscape quickly became the contempo- 
rary source for information on local natural environ- 
ment issues. It has provided an ideal bridge between 
the recording of natural environment information for 
its own sake and in directing and assessing conserva- 
tion priorities (Brunton 1986b). To some, the timeliness 
and thus effectiveness appears to have suffered, how- 
ever, after a decision to reduce the publication sched- 
ule, for manpower and economic reasons, from five 
issues/year to four, starting with the 1988 volume. 
The subsequently reduced publication frequency and 
fewer total pages have coincided with a substantial 
reduction (by almost half) in the number of articles 
with direct conservation applications. 

The huge increase in membership and higher profile 
of conservation and field-oriented activities in the late 
1960s may have set too hectic a pace for some long- 
time Club stalwarts who had been more concerned with 
the national publication role. Hoyes Lloyd resigned 
from the Council at the end of 1967, stating that he 
felt it inappropriate as an OFNC Honorary Member 
to also serve as a Club director (LAC OFNC Collec- 
tion, 6 June 1967). There is no such limitation on Hon- 
orary Members, however, and a number have contin- 
ued to serve on Council with distinction for many years. 
Notable in this regard is 1979 Honorary Member Bill 
Cody, who has served continuously on Council since 
December 1947, shortly after becoming The CFN 
Business Manager earlier that year (LAC OFNC Col- 
lection, 21 May 1947). And similarly impressive is the 
contribution of Frank Pope, who has served on Council 
continuously since 1980 and has occupied the positions 
of Corresponding Secretary (three years), Treasurer 
(six years), Vice-President (one year) and President 
(an unprecedented six years) (Appendix 1). 

The Canadian Field-Naturalist underwent a signif- 
icant reconfiguration in January 1970, introducing a 
new size, format and appearance not just to improve 
its look but “to [make] its content as relevant as pos- 
sible to the natural history needs of our time” (Mosquin 
1970). And by “relevant”, Editor Ted Mosquin meant 
strongly conservation oriented. That first issue was 
graced by a photo with a Timber Wolf — the contem- 
porary symbol of wilderness protection — crossing a 
snowy landscape. The issue contained both a variety 
of articles on Canadian endangered species and a direc- 
tory of conservation organizations in Canada. Conser- 
vation applications of carefully-researched ecological 
and natural diversity investigations have been a hall- 
mark of The CEFN ever since. 


Vol. 118 


Both as OFNC 1969-1971 President and editor of 
the CFN from 1967 to 1972, Mosquin was at the fore- 
front of efforts to enhance the level of OFNC conser- 
vation action and to engage both the membership and 
the general public in this. OFNC 1971-1972 President 
Sheila Thomson and OFNC 1972-1975 President 
Irwin Brodo were similarly motivated conservation 
voices in the OFNC, the three of them working togeth- 
er to persuasively direct Club environmental protection 
initiatives. Mosquin initiated a series of at times hard- 
hitting and at times controversial editorials (not the 
last we were to see in the CFN!), challenging Club 
members, the larger naturalist community, and public 
decision-makers to be more effective in protecting 
natural environments in Canada. In 1972 he resigned 
the CFN editorship to take on the challenge of found- 
ing editor of the Canadian Nature Federation’s Nature 
Canada. 

The infectious enthusiasm, environmental passion 
and unquestioned technical expertise of these three 
activists in concert with other Club leaders of the day 
such as OFNC 1963-1966 President George McGee 
(1909-1991) and OFNC 1967-1969 President Hue 
McKenzie, were an inspiration to a whole generation 
of new Ottawa-area naturalists (personal observation). 
For us newcomers, they transformed the understand- 
ing of the study of the natural environment from a 
seemingly slightly idiosyncratic, individual endeavour 
undertaken for personal entertainment to a meaning- 
ful — even important — and highly integrated calling. 
What a revelation! 

The growing commitment to natural environment 
conservation in the late 1960s and early 1970s saw 
the re-energizing of OFNC Council and membership 
support in this area. At the urging of Ted Mosquin and 
Ed Greenwood and recognizing that the Preservation 
of Natural History Sites Committee was not broad 
enough in its mandate, in early 1967 the OFNC estab- 
lished the Natural Areas Committee to coordinate 
original field conservation research and to actively 
search out potential natural reserve sites in the Ottawa 
Valley (LAC OFNC Collection, 10 April 1967). By 
1970, the committee was undertaking a broad program 
of local field investigations and had provided a basis 
for the identification of significant natural areas in 
the Regional Municipality of Ottawa-Carleton’s 1974 
Official Plan. So significant was the role and mandate 
of this group that the Natural Areas Committee was 
soon being promoted as the most significant of Club 
committees (LAC OFNC Collection, 8 December 
1970). Increasing demand for input into various con- 
servation issues both locally and beyond, resulted in 
the establishment of the Research and Briefs Commit- 
tee (1972-1973), which in 1974 was combined with the 
Natural Areas Committee to form the Conservation 
Committee. This committee continues to play an inte- 
gral role in OFNC and community affairs to the 
present day. 


2004 


The OFNC Centennial in 1979 understandably 
brought forth a great deal of interest in the history of 
the Club in particular and of the naturalist commu- 
nity of Ottawa in general. Trail & Landscape provided 
the major vehicle for sharing this information amongst 
the local membership. Events culminated with a ban- 
quet intended to be on the exact 100" anniversary, 
but as with previous celebrations, it was erroneously 
held on 19 March (Reddoch 1979a), not the actual 
centennial date of 25 March. 

The increased interest in the historical record gen- 
erated by the Centennial year encouraged a new appre- 
ciation of the designation of OFNC Honorary Mem- 
bers. These had been given out only irregularly over 
the years to honour lengthy, significant service to the 
OFNC or Canadian natural sciences. Although honor- 
ary memberships have been awarded more regularly 
since 1971, the Centennial Year saw a new focus on 
this acknowledgment of such exceptional contributions 
(Brunton and Gummer 1987). 

The conservation initiatives of the 1970s and early 
1980s saw a resurgence of more systematic field 
examinations of uncommon and ecologically significant 
habitats in the Ottawa Valley such as fens, rivershore 
communities, and relict woodlands (Dugal 1978; 
Reddoch 1979b; White 1979). Publication of the first 
complete, annotated checklist of the vascular plants 


BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 25 


of the Ottawa District also reflected this knowledge 
(Gillett and White 1978) and contained many of the 
records newly discovered by Club members. 

That burst of new knowledge and energy demon- 
strated that suggestions in the early years of the 20" 
century that our knowledge of the natural environ- 
ment of the Ottawa area was all but complete, were 
badly incorrect. Our understanding of native biodi- 
versity and of the ecological systems supporting it 
requires continual inventory and evaluation to enable 
effective protection and maintenance of this natural 
heritage. 

The Club’s conservation activities in this era, led 
by activist presidents Roger Foxall and Roger Taylor 
between 1978 and 1982 and by such energetic mem- 
bers as Loney Dickson, Allan Reddoch, Joyce Red- 
doch, Albert Dugal, Stephen Darbyshire, and David 
White, covered an amazing spectrum and exhausting 
number of subjects. These ranged from a campaign for 
a conservation area system in the Region of Ottawa- 
Carleton (now city of Ottawa) to prevention of the 
importation of Raccoon-dogs into Canada, and from 
the prevention of 1988 Winter Olympic facility devel- 
opment within Banff National Parks to consultations 
with federal government officials on ecological re- 
quirements in Gatineau Park and National Capital 
Greenbelt planning. 


Ficure 16. Excursion of Ottawa Field-Naturalists to the Rideau Trail in October 1979 (photograph by C. Beddoe). Contrast 
with Figure 9. Long-time member Mary Stuart is second from left. 


26 THE CANADIAN FIELD-NATURALIST 


The Club hosted the annual meeting of the Feder- 
ation of Ontario Naturalists in 1983, establishing a 
successful program format that was followed by sub- 
sequent provincial gatherings for years to come. The 
OFNC again successfully hosted the provincial orga- 
nization’s annual conference in 1993 (Pope 1993). 

A more whimsical contribution is to wildlife con- 
servation initiated in this period was the ““Seed-a-thon’”’, 
a fund-raising effort whereby sponsors pledge a par- 
ticular amount for every species observed on a par- 
ticular day by a particular birding team, to be applied 
towards the cost of supporting a network of OFNC 
public bird feeder stations. Initiated by the Birds Com- 
mittee on the suggestion of long-time OFNC member 
Violet Humphries in 1981 (personal communication), 
this was one of the first of what now are commonplace 
fund-raising events across Canada (Brunton 1981). 

There was a human cost to all this activity, how- 
ever, and many of the lead Club personnel in these 
matters eventually were called away by the mundane 
but critical matters of family and careers, moving on 
to other things and/or were simply exhausted by it all. 
This was exacerbated in the later years of the 1970s 
and into the 1980s by new pressures on the time and 
resources of professional research staff within the 
federal public service, reducing their availability and 
energy for involvement in such initiatives as local 
conservation and research. Only three OFNC presidents 
since the mid 1970s (D. F. Brunton, R. John and E. 
Zurbrigg), for example, were professionally involved 
in natural environment investigation or management 
(Appendix 1). 

One of the most satisfying conservation ventures 
of recent decades has to be the effort begun in the 
early 1980s to save the massive, Provincially Signifi- 
cant Alfred Bog east of Ottawa (Cuddy 1983) from 
destruction by agri-business interests. This battle saw 
the Club purchase conservation land for the first time 
(in 1982) so as to have legal standing in the fight to 
secure proper protective zoning. It was a long and 
difficult effort but with the effective leadership of 
OFNC President Frank Pope, considerable financial 
and material contributions from numerous individual 
members, and the critical involvement of the Nature 
Conservancy of Canada, the entire wetland was saved 
and is now either in protective public ownership or 
zoned as parkland (Pope 2002). 

The primary national contribution of the OFNC 
continued to be led by the publication of the CEFN, al- 
though as noted, direct involvement in national-scale 
conservation issues was common enough in this peri- 
od. Despite a continuing and perhaps even enhanced 
national/international status and production standard, 
however, the seemingly unanswerable question of how 
national or local the journal should be was raised again 
in a series of editorials in the late 1970s. The appro- 
priateness and credentials of a citizen-based (‘‘ama- 
teur”) organization directing the affairs of the journal 
was severely criticized by Lorraine Smith, Editor 1972- 


Vol. 118 


1981, who suggested that “the burden of being account- 
able to the scientific natural history community for a 
national journal ... may now be more than this local 
club should retain.” [The days of calling the OFNC a 
“scientific society” were truly gone!] “Therefore... if 
the journal’s reputation is not maintained and there is 
no move for a change in publisher, I challenge those 
who want a top-quality Canadian natural history jour- 
nal, such as The Canadian Field-Naturalist is now, to 
initiate a new journal” (Smith 1981). These comments 
do not explain, however, how the OFNC managed to 
maintain and enhance such a fine journal for so many 
years. 

After much discussion, consultation and review by 
an Ad Hoc Committee on OFNC Publications com- 
posed primarily of scientific specialists, the OFNC 
Council approved (8 December 1982) a new Publica- 
tions Policy to guide the CEN and other Club publi- 
cations (Bedford 1983). The Policy reiterated the Club’s 
continuing, century-long commitment to the scientific 
integrity and high technical standards of the CFN, as 
well as the Club’s intention to have the journal remain 
the official publication of the OFNC. 

Francis Cook assumed editorship of the CFN from 
1962 to 1966, and again in 1981, and has guided the 
journal to the present day with a steady and depend- 
able hand, introducing important innovations such as 
the publication of Canadian status reports for candidate 
endangered species. There has been a great increase 
in the amount of material published in each volume 
during Cook’s tenure (volumes averaged 756 pages 
over the five years ending in 2002). In the face of the 
many complications in the publication process, the 
timely publication of some issues has not occurred. 
Scientific and technical quality has never been sacri- 
ficed, however. The 28 volumes edited by Cook repre- 
sent by far the longest and most prolific service of any 
CEN editor in the 124-year history of the publication, 
exceeding by more than a decade the tenure of the 
next longest-serving editors, Harold Senn and Arthur 
Gibson (Brunton 1986a). 


Contemporary Times and the Future (mid- 
1980s to date) 

James Macoun, Percy Taverner and other “nation- 
als” who worked so hard to enhance the mandate, 
effectiveness, and reach of The Canadian Field- 
Naturalist would be relieved to see that the publica- 
tion has achieved the solid reputation, steady produc- 
tion, and high scientific standards they hoped to see. 
An interesting expression of that long-term contribu- 
tion and durability is demonstrated in the number of 
original descriptions (diagnoses of new taxa) which 
have occurred in the pages of the CFN. Over 730 have 
been published in the first 100 volumes (Brunton 
1987). 

The level of field-oriented investigation of the Otta- 
wa Valley natural environment that was so productive 
in the 1970s and early 1980s, however, has been dras- 


2004 


BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 


Number of Members 


| SS ES SSS CY MSY ET | 
TS 


prerereeco 


FiGureE 17. Ottawa Field-Naturalists’ Club membership trend 1975-2003. Numbers of institutional subscribers to The Canadian 
Field-Naturalist were not calculated within membership totals nor reported separately in Annual Reports after 1971. 
A 1975 reorganization offered individual members (reported to total 1371 that year) the opportunity to be only non- 
voting subscribers to the The Canadian Field-Naturalist, dramatically reducing the OFNC membership total reported 
for 1976 (Erskine 1977) [Individual and institutional subscribers were reported separately in the Annual and Editor’s 
Reports for The Canadian Field-Naturalist in each volume]. Absolute membership numbers prior to 1976, therefore, 
are not directly comparable to these of subsequent years (from OFNC Annual Reports, 1975-2002). 


tically reduced. The appearance of substantially fewer 
area inventories and record documentation articles in 
Trail & Landscape reflects that trend. This may be part 
of an unfortunately strong North American perception 
in recent decades that continuing, original field in- 
vestigations are of less importance than in previous 
times. T&L remains, however, an important and highly 
relevant source of environmental news and natural 
environment documentation. 

Programs intended to actively involve the partici- 
pation of local members have remained an important 
part of the OFNC. Dozens of field outings, lectures 
and special events are undertaken each year involving 
a diversity of well-informed leaders and speakers. 
For decades the Club’s Excursions and Lectures Com- 
mittee has crafted an interesting and instructive program 
that contributes greatly to the enhancement of members’ 
skills and to their enjoyment in the field. While field 
investigations focus on both familiar and exotic local- 
ities throughout the Ottawa District and the Ottawa 
Valley, the Club does not confine itself to this area. 
The spring “migration” of a bus load of OFNC mem- 
bers to Point Pelee National Park, for example, has 
become a popular biennial birding event since the 1980s. 


This is important stuff. The majority of Club mem- 
bers are locally based and most of us are involved, 
first and foremost, because we enjoy exploring and 
learning about our natural world. Providing support 
for scientifically important undertakings such as the 
publication of the CFN and the protection and preser- 
vation of important natural landscapes, is a valuable 
additional membership benefit. Membership in the 
OFNC remains an easy and enjoyable way for indi- 
vidual citizens to support greater understanding of, 
and protection for, important natural landscapes and 
features in the Ottawa Valley and beyond. 

A noticeable change in the role of the local Club in 
regards to its community has occurred in recent years. 
The OFNC is much more integrated into environmen- 
tal decision-making at a municipal and provincial 
level than it has been at any time in its past. It has 
become, in an sense, an ecological consultant to that 
community. While such a role requires seemingly end- 
less attendance at meetings and reduced time for core 
field work, it has provided many excellent opportunities 
for Club representatives to bring accurate ecological 
information into the discussion of an issue at an early 
and effective stage of the decision-making process. A 


28 THE CANADIAN FIELD-NATURALIST 


cynic might say that the Club has become somewhat 
bureaucratized; an optimist would suggest that the Club 
now more frequently has a seat at decision-making 
tables across the National Capital Region. It is impor- 
tant to note, for example, that it was through this per- 
iod that the final protection of the important Alfred 
Bog natural area was achieved. 

Another very positive achievement of the last decade 
was the establishment of the Louise de Kiriline Law- 
rence Conservation Action Fund in 1994, founded with 
a generous bequest from the estate of that outstanding 
and eloquent field naturalist, Louise de Kiriline Law- 
rence (1894-1992) (Ainley 1994). It is sustained by 
further donations and by the profit from the sale of 
particular OFNC products such as the “Natural Areas” 
book (Brunton 1988). The fund provides for “stra- 
tegic and timely expenditures towards the conservation 
of natural areas” in the Ottawa Valley (Anonymous 
1994). 

A continuing interest for many Club members is 
the Fletcher Wildlife Garden (FWG), established in 
1987 in degraded woodland and regenerating agricul- 
tural land on National Arboretum property near the 
Rideau Canal. With the active cooperation and assis- 
tance of Central Experimental Farm (Agriculture 
Canada) personnel who are responsible for land man- 
agement here, the Fletcher Wildlife Garden volun- 
teers have transformed the site with native plantings, 
the creation of a pond and through public interpre- 
tation. The objective is not only to produce on-going, 
evolving habitat restoration but to demonstrate suita- 
ble wildlife-friendly landscaping and gardening prac- 
tices for National Capital residents and visitors alike 
— to be “a model for urban gardeners” (Conservation 
Committee 1987). 

These are all positive features. A disturbing trend in 
Club affairs, however, has been declining membership 
over the last two decades (Figure 17). Perhaps not 
coincidentally, Club membership peaked ca. 1982 to 
1985, at the end of the period of greatest conservation- 
oriented activity. The local membership decline is 
seemingly in tandem with the decline of original field- 
based investigations. Nonetheless, the number of 
OFNC members remains substantial — the Club is still 
probably the largest regional natural history organi- 
zation in Canada — but the downward membership 
trend needs reversing (Figure 17). 

The reasons for this decline in participation are un- 
clear. It seems unlikely to be attributable to the ready 
availability of up-to-date, comprehensive natural en- 
vironment data on the World Wide Web since the mem- 
bership decline began well before such technology 
was widely available. It may, in part, be a function of 
some wider, societal disinterest in hands-on involve- 
ment with citizen-based, research-oriented groups. 
Whatever the reason(s), the decline has been substan- 
tial and if it continues, will undermine the financial 
security and effectiveness of the OFNC. 


Vol. 118 


Since the 1990s the digital world has had as dra- 
matic an impact on the OFNC as it has on so many 
facets of contemporary Canadian life. We are only 
just coming to terms with its potential benefits and 
challenges. The OFNC’s impressive web page (http:// 
www.ofnc.ca) has allowed for rapid internal commu- 
nications amongst Club officials, committee members 
and conservation partners, and for the posting of 
information and timely news items. Digital commu- 
nication is only the latest expression of the Club’s 
commitment to education of our membership and of 
the general public in regard to important natural envi- 
ronment features, needs and opportunities. The impor- 
tance of the Club’s educational programs has been a 
major theme throughout OFNC history and is ex- 
pressed in the mission statement of the organization 
(see frontispiece of this issue). There is every indica- 
tion that it will remain so into the future. 

It is not yet clear how the Club will move forward 
from this foundation of established programs and re- 
sources in order to stay informed, relevant — and sol- 
vent — into the future. That’s not a new challenge but 
is one that must be answered by each generation in 
response to the circumstances and opportunities of 
their day. 

Some things are clear. For one, the Club needs to 
re-establish a higher level of field-oriented investigation 
amongst its membership. This will not only maintain 
and develop the skills and interests of present mem- 
bers but also inspire and generate new members. Ac- 
curate and timely field investigation and documen- 
tation are essential foundations for the maintenance 
of the high level of technical credibility that the Club 
has earned over the years. Continuing to do our home- 
work, getting the facts straight, and presenting such 
information in a defensible, objective manner are the 
keys to maintaining and enhancing the OFNC’s impre- 
ssive record of conservation achievement. 

As with any organization, regardless of how vener- 
able, the OFNC must also continue to re-examine what 
it provides for the individual member. This is perhaps 
even more critical in an era with an unprecedented 
number of electronic diversions competing for the 
attention of both members and partners alike. Will the 
traditional superb program of excursions, lectures, and 
publications be enough to inspire and encourage a 
growing, involved membership? Do the new technol- 
ogies of our time offer the same manner of new op- 
portunities that technologies like the railway train, 
electric lighting, the automobile, and the airplane 
offered earlier OFNC members? They surely do. 

What I hope is a clear message from this review of 
the long, remarkably productive journey of Ottawa 
naturalists is that the OFNC is not a static, hard-and- 
fast “thing”. It is a community, an association of like 
minds that aims to explore and celebrate the magnifi- 
cent natural world of the Ottawa Valley and of Canada. 
And as with any community, we in the OFNC have 


2004 


BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 29 


tos 


FIGURE 18. Record service to The Canadian Field-Naturalist: Left: Francis R. Cook, Editor 1962-1966; Associate Editor 
(Herpetology) 1972-1981; Editor 1981-present; right: William J. Cody, Business Manager 1948-present. 25 August 


2004. Photo courtesy Ron Bedford. 


and have had the benefit of the skills, insight, and 
inspirational leadership of numerous individuals and 
groups over the years. 

In weak times and strong, the common thread that 
has bound Ottawa naturalists since 1879 — indeed, since 
1863 — is an overwhelming desire to contribute to the 
protection and enhancement of the natural features 
and values that make this place so special. At the end 
of the day it comes down to the fact that James Fletcher, 
Frank Pope, Percy Taverner, Sheila Thomson, Edward 
Van Cortlandt, Herb Groh, Gordon Hewitt, and the rest 
of us are not that different ... we’re all just Ottawa 
field naturalists. 


Acknowledgments 

The daunting challenge of reviewing a manuscript 
of this nature was undertaken in a timely and helpful 
manner by a team of naturalists well familiar with the 
OFNC and OFNC personalities. These reviewers were 
Ron Bedford, Bill Cody, Francis Cook, Karen McIntosh, 
Elizabeth Morton, Frank Pope and Joyce Reddoch. 
They added much to the effort and I gratefully ack- 
nowledge their input. I remain, however, fully respon- 
sible for any errors or omissions which may have 
escaped their sharp eyes. Frank Pope and Francis 
Cook provided additional factual input throughout 
the development of the manuscript. This review has 
also benefitted from the innumerable discussions (and 


debates!) I have enjoyed with Francis Cook over the 
last 25 years on numerous subjects related to the his- 
tory of the OFNC and to the study of Canadian natural 
sciences. Ron Bedford, Frank Pope, Joyce Reddoch, 
and Rob Lee (present chair of the Macoun Field Club) 
were instrumental in obtaining many of the illustra- 
tions employed in this article. Rob Lee and Joyce Cook 
scanned several of these. I am happy too, to acknowl- 
edge two other important players in this effort. First 
is the late Herb Groh whose clipping file on OFNC 
personalities and whose interest in the Club’s past 
inspired my own curiosity. The other gratefully ack- 
nowledged is the staff at the Library and Archives 
Canada (formerly the National Archives of Canada) 
who so efficiently and pleasantly facilitated my 
research with the OFNC collection and related files 
held within the walls of that national treasure. 


Documents Cited (marked * in text) 

Dorais, M., R. Bouffard, R. Chenier and K. Adamson. 
1974. An examination of the past and present status of the 
Mer Bleue Peat Bog near Ottawa, Ontario. [Unpublished 
report], Faculty of Science and Engineering, University 
of Ottawa. 105 pages. 

LAC OFNC Collection. Ottawa Field-Naturalists’ Club 
Collection (MG 28 I 31), Library and Archives Canada, 
Ottawa. 

LAC Taverner Collection. Percy A. Taverner collection, 
Library and Archives Canada, Ottawa. 


30 THE CANADIAN FIELD-NATURALIST 


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32. THE CANADIAN FIELD-NATURALIST 


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Received 22 June 2004 
Accepted 10 August 2004 


END NOTES: 

' Number 1 (un-numbered), Braddish Billings’ Ottawa 

vascular plant list (Billings 1868) was published between 

March and November 1868; Number 2 (numbered), 500 

copies of Sir J. A. Grant’s examination of the surficial 

geology of the Ottawa Valley (Grant 1868) were published 

December 1868; Number 3 (numbered), Thomas Wily’s 

discussions of swallows (Wily 1869) was published De- 

cember 1869 (LAC OFNC Collection, 10 December 1869; 

Dore 1968). 

Much later, Henry Ami reminiscing about events leading 

up to the formation of the Club described *... how the 

Ottawa Club originated from the Epping Forest and Field 

Club, and later Ottawa Naturalists Club ...” (LAC OFNC 

Collection, 2 February 1924). Ami may, however, have 

been referring to names applied to informal groupings of 

field associates, such as the Fletcher, Harrington et al. 
group which conducted field work together in the Billings 

Bridge area in the late 1870s (Harrington 1909). There is 

no further mention of these groups in OFNC records by 

any of the individuals who, unlike Ami, were actively 
involved in the formation of the Club. 

Later, R. B. Whyte erroneously reported this as occurring on 

19 March 1879 (Whyte 1880). He may have been referring 

to an earlier preliminary meeting or to the date of the 

OLSS Council meeting where Fletcher and Henry Small Jr. 

obtained an expression of the co-operation and assistance 

from the parent organization, rather than the formal orga- 
nizational gathering. Being documented only one year after 
the fact and by a Club founder and the originator of the 
idea of the OFNC, this error would quite understandably 
be accepted as fact and was repeated for over 100 years 

as the formal starting date of the OFNC (e.g. Groh 1955; 

Taylor 1979). 

+ Relations between The Ottawa Field-Naturalists’ Club and 
The Ottawa Literary and Scientific Society appear to have 
been strained on at least several occasions in the following 
years. In his 30 November 1886 Inaugural Address, for 
example, OLSS President W. P. Anderson complained 


nN 


Vol. 118 


that the OFNC (of which he was a founding member and 
had been the Treasurer for the previous three years!) was 
“antagonistic to the Ottawa Literary and Scientific Soci- 
ety” (LAC OFNC Collection, 3 December 1886). The 
OFNC and OLSS councils individually discussed the prob- 
lem, the OLSS Council subsequently stating that they 
would consider “ ... any proposition which the field Club 
may wish to make looking for a change in the relations 
between the Club and the Society but that a joint meeting 
of the two Councils cannot conveniently be arranged ...” 
(LAC OFNC Collection, 14 January 1887). Although no 
such joint meeting or other formal Club action resulted, 
the OFNC Council continued to meet in the OLSS 
museum until March 1890. That was despite another 
minor dust-up in 1889 concerning the OFNC’s offer to 
cover some of the public lecture programming required 
of the OLSS in order to maintain its Ontario government 
grant (LAC OFNC Collection, 6 November 1889). 

The involvement of women in the OFNC was very much 
promoted in these early years — in a patronizing if sincere 
way. The launch of The Ottawa Naturalist was accompa- 
nied by a special plea for the involvement of women ... 
“especially will be ladies welcomed to our ranks, and every 
effort will be put forward to make the excursions and 
soirees pleasant as well as instructive” (Harrington 1887). 
At the March 1888 Annual Meeting “... a discussion en- 
sued as to the eligibility of ladies as officers, and the 
desireability of having some on the Council. Professor 
Macoun gave notice that he would at the next General 
meeting of the Club move that the executive Committee 
consist of six members, three of whom shall be ladies” 
(LAC OFNC Collection, 20 March 1888). This was accom- 
plished by a Constitutional amendment in March 1890. 
Margaret A. Mills became the first female OFNC Officer 
when “Miss Mills” was elected Second Vice-President in 
1892 (LAC OFNC Collection, 15 March 1892). She was 
not present at the meeting, however, and was apparently 
not agreeable to her election, as she resigned effective the 
next Council meeting 9 days later (LAC OFNC Collection, 
24 March 1892), but stayed on as a Club member for 
several more years. 

“This [Ottawa] District was then [1880] understood to 
mean a radius of about twelve miles from the City of 
Ottawa ... latterly, however, with the general consent of 
the botanists of the club, this radius has been extended to 
about 30 miles ...” (Fletcher 1888). The Ottawa District 
boundary was metricated in 1981 to a 50 km radius circle 
centred on the Peace Tower of the Parliament Buildings 
(Anonymous 1981), including landscape within both south- 
eastern Ontario and southwestern Quebec. 

The portrait hung for many years in the Geological Survey 
Museum (later, National Museum of Canada). It now can 
be seen on display in the Logan Gallery of the Geological 
Survey of Canada headquarters on Booth Street in Ottawa. 
The saga of the OFNC library does not end there, however. 
In 1948 the Ottawa Public Library (OPL) reported to the 
Club that they had discovered “... the old library of the 
Club, now deposited in a storeroom adjacent to the main 
library” and that several hundred dollars worth of Canadian 
Field-Naturalist (CFN) back numbers were recovered 
(LAC OFNC Collection, 29 October 1948). The Council 
agreed to thank the OPL for storing the collection (since 
1917!) by donating a set of The CFN back to 1935. 
Although the OPL apparently offered to continue to store 


2004 


a 
oO 


the 24 shelves worth of material at no cost to the Club, 
Council determined that all of the natural history and 
geology books should be sold (LAC OFNC Collection, 
22 November 1948). No one was aware, apparently, that 
the Club had given the library to the OPL back in 1917 and 
thus was obviously in no legitimate position to sell the 
material. Nonetheless, the library was sold to “Mr. 
[Bernard] Amtmann, a dealer in second hand books ...” 
who assured the Council that OFNC members “would be 
given first choice before the [collection] catalogues were 
sent out ... through his extensive mailing list” (LAC OFNC 
Collection, 19 January 1950). The Club’s initial share of 
sales before Amtmann moved to Montreal with the collec- 
tion was $169 (LAC OFNC Collection, 20 November 1950); 
the following year Amtmann offered the Club $200 for the 
remaining volumes of the library which he had sold off 
“on a poundage basis” and for which he claimed to have 
incurred a considerable loss. The offer was accepted (LAC 
OFNC Collection, 16 November 1951) ... a sad ending 
to what must have been a magnificent collection with an 
intriguing provenance. 

The portrait by Franklin Brownell was commissioned in 
late 1911 (LAC OFNC Collection, 18 December 1911) 
and hung in “a suitable and prominent place” in the Ottawa 
Public Library in the spring of 1912 (LAC OFNC Collec- 
tion, 12 March 1912). In the 1920s it was transferred to 
the National Museum of Canada “for safekeeping” (LAC 
OFNC Collection, 14 December 1926) where it hung for 
many years in the herbarium, its actual ownership and 
the “temporary” nature of its placement at The Museum 
eventually forgotten. When this history was discovered and 
brought to the attention of National Museum of Natural 
Sciences Assistant Director (and former OFNC Vice-Pres- 
ident) C. G. Gruchy and Agriculture Canada staff in 1985 
(personal communication), arrangements were made by 
The Museum for cleaning and minor restorations to be 
undertaken. The portrait, still technically owned by the 
Club and on loan to The Museum, was officially donated to 
The Farm on 2 June 1986 by OFNC 1986-1988 President 
W. G. Gummer (1915-1999) as part of the celebrations of 
100 years of agricultural research in Canada (Hall 1986). 
The Museum and the Club had now fully honoured their 
commitment to the memory of James Fletcher. The portrait 
presently hangs, appropriately enough, in the William 
Saunders Building, named after the Central Experimental 
Farm Director who hired Fletcher as Canada’s first Domin- 
ion Entomologist and Botanist. 

The Ottawa Naturalist was published quarterly for a brief 
period (1889/1890) as a cost-saving measure but reverted 
to monthly publication (at the urging of Will Harrington) 
because of reader preference for more frequent produc- 


BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 


w 


15 


33 


tion. This was initiated despite the Club’s request for a 
provincial publication assistance grant being denied. An 
avowed Tory, John Macoun proclaimed that “‘politics had 
intervened with the success of the application and that we 
would have to learn to vote properly before we could hope 
to succeed ...” (LAC OFNC Collection, 18 March 1890). 
The none too subtle shot at the long serving Liberal govy- 
ernment of Oliver Mowat was quite likely an accurate 
observation of the politics of that day (Gywn 1984). The 
Ottawa Naturalist did eventually receive a provincial gov- 
ernment publication grant (of $300.00), beginning in 1897. 
But presumably to Macoun’s chagrin, this occurred with 
the Liberals still in power! Provincial financial assistance 
was received thereafter until 1924 (LAC OFNC Collection, 
3 November 1924), as was generous federal publication 
support funding for The Canadian Field-Naturalist in the 
1980s and 1990s. 

The Macoun Autobiography was reprinted as an OFNC 
Special Publication in the Club’s centennial year (Macoun 
1979), with an new introduction by Richard Glover and 
with both editorial notes and a biographical sketch by 
historian William A. Waiser. 

P. B. Symes presented his “nearly complete set” of The 
Canadian Field-Naturalist and its predecessors to the 
Club in 1923. It formed the basis for “the Club’s set of 
original issues” (Patch 1923) which is maintained for the 
purposes of the current CFN editor. The set is now complete 
(Francis Cook, personal communication). 

When Pauline Snure was elected in 1948, she was the first 
woman president of the OFNC, despite the conspicuous 
affirmative action efforts on behalf of women members by 
James Fletcher and associates in the 1880s and again in 
the first decade of the 20" century. 

It was presumably this 75" anniversary that inspired Herb 
Groh to develop a biographical clipping file on OFNC 
Presidents and other officers. The file was presented to the 
Club in the early 1980s by Pierre Taschereau on behalf of 
Groh’s widow and is now preserved in the LAC OFNC 
Collection. 

The present editor has suggested (personal communication) 
that this treatment might be overly critical of Senn’s edi- 
torial tardiness in comparison to the at-times substantial 
delay of issues of the CFN during his (Cook’s) tenure. 
While recent delays sometimes approached (though 
never matched) those of the 1950s, the critical difference 
between the two was the impact on the Club in their res- 
pective eras. The substantial publication delay in the early 
1950s all but shut down Club productivity. While not to 
suggest that contemporary publications delays are without 
impact, they are clearly less threatening than those of 50 
years ago. 


34 


THE CANADIAN FIELD-NATURALIST 


APPENDIX 1: Ottawa Field-Naturalists’ Club officers (1879-2004) 


(derived from (LAC OFNC Collection Minute Books (1879-1970) and The Canadian Field-Naturalist listings and contem- 


porary OFNC Council Minutes) 


1879/1880 


1880/1881 


1881/1882 


1882/1883 


1883/1884 


1884/1885 


1885/1886 


1886/1887 


1887/1888 


1888/1889 


1889/1890 


1890/1891 


1891/1892 


1892/1893 


1893/1894 


1894/1895 


1895/1896 


1896/1897 


1897/1898 


1898/1899 


1899/1900 


1900/1901 


1901/1902 


1902/1903 


1903/1904 


1904/1905 


1905/1906 


1906/1907 


President 


W. White 


J. Fletcher 

W. R. Riddell* 
J. Fletcher 

J. Fletcher 

H. B. Small Jr. 
H. B. Small Jr. 
W. H. Harrington 
John Macoun 
R. B. Whyte 
R. B. Whyte 
R. W. Ells 

R. W. Ells 


R. W. Ells 


G. W. Dawson 


G. W. Dawson 
G. W. Dawson 
F. Shutt 

F. Shutt 

EWE. Prince 
BE Prince 
H. M. Ami 

H. M. Ami 

R. Bell 

R. Bell 

W. T. Macoun 
W. T. Macoun 
S. B. Sinclair 


W. J. Wilson 


Vice-President(s)! 


J. Fletcher 

W. P. Riddell* 
W. D. LeSueur* 
R. B. Whyte 
W. D. LeSueur 
R. B. Whyte 
H. B. Small Jr. 
R. B. Whyte 

J. F Whiteaves 
R. B. Whyte 

F. R. Latchford 
R. B. Whyte 

J. Fletcher 
John Macoun 
S. Woods 

R. B. Whyte 

S. Woods 

John Macoun 
S. Woods* 

C. F. Marsan 
R. W. Ells 

H. M. Ami 

J. Ballantyne 
R. B. Whyte 

J. Ballantyne 
H. M. Ami 

T. S. MacLaughlin 
W. H. Harrington* 
M. A. Mills* 
F. Shutt 

R. W. Ells 

F. Shutt 

J. Fletcher 

F. Shutt 

H. M. Ami 


John Macoun 
W. S. O'Dell 
R. Bell 

D. A. Campbell 
W. T. Macoun 
A. E. Attwood 
W. T. Macoun 
A. E. Attwood* 
A. Halkett 

W. J. Wilson 

S. B. Sinclair 
W. J. Wilson 

F. Shutt 

F. Shutt 


Treasurer 


R. B. Whyte? 


W. H. Harrington? 
W. H. Harrington? 
W. P. Anderson 
W. P. Anderson 

W. P. Anderson 

T. S. MacLaughlin 
T. S. MacLaughlin 
J. Fletcher 

J. Fletcher 

J. Fletcher 

J. Fletcher 

G. Harmer 


A. G. Kingston 


A. G. Kingston 


.Am 
"Harrington 
ig* 


J. Fletcher 
J. Fletcher 
J. Fletcher 
J. Fletcher 
A. Gibson 
A. Gibson 
A. Gibson 
A. Gibson 


A. Gibson 


Vol. 118 


Secretary(ies)” 
R. B. Whyte? 


W. H. Harrington? 
W. H. Harrington? 
W. H. Harrington 
W. H. Harrington 
W. H. Harrington 
W. P. Anderson 
W. H. Harrington 
W. H. Harrington 
T. S. MacLaughlin 
T.S. MacLaughlin 
T. S. MacLaughlin 
W. H. Harrington 


H. M. Ami 


H. M. Ami 
H. M. Ami 
A. Halkett 

A. Halkett 

A. Halkett 
W. J. Wilson 
W. J. Wilson 
W. J. Wilson 
W. J. Wilson* 
W. J. Wilson 
W. J. Wilson* 
T. E. Clarke 
T. E. Clarke 


T. E. Clarke 


2004 


1907/1908 
1908/1909 
1909/1910 
1910/1911 
1911/1912 
1912/1913 
1913/1914 
1914/1915 
1915/1916 
1916/1917 
1917/1918 
1918/1919 
19193 
1920 

1921 

1922 

1923 

1924 


1925 


1926 
1927 
1928 


1929 


1930 
1931 
1932 
1933 
1934 
1935 


1936 


BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 


President 


W. J. Wilson 

A. E. Attwood 
A. E. Attwood 
A. Halkett* 

A. G. Kingston* 
A. McNeill 

L. H. Newman 
L. H. Newman 
A. Gibson 

A. Gibson 

H. I. Smith 

H. I. Smith 

C. G. Hewitt 
M. Y. Williams 
M. Y. Williams 
R. M. Anderson 
R. M. Anderson 
H. Lloyd 

H. Lloyd 


H. Lloyd* 
G. A. Miller* 


N. Criddle 
N. Criddle 
E. M. Kindle 


E. EF. G. White 


H. F. Lewis 

H. F. Lewis 

C. M. Sternberg 
C. M. Sternberg 
M. E. Wilson 
M. E. Wilson 


H. Groh 


Vice-President(s)! 


A. E. Attwood 
A. E. Attwood 
A. Halkett 

A. Halkett 

E. FG. Eifrig 
A. Halkett 

E. E.G. Eifrig 
A. G. Kingston 
L. H. Newman 
A. Gibson 

L. H. Newman 
A. Gibson 

J. W. Gibson 
A. Gibson 

H. I. Smith 

C. G. Hewitt 
H. I. Smith 

C. G. Hewitt 
H. I. Smith 

C. G. Hewitt 
E. D. Eddy 

M. Y. Williams 
C. G. Hewitt 
L. D. Burling 
M. Y. Williams 
L. D. Burling 
P. A. Taverner 
L. D. Burling 
R. M. Anderson 
G. A. Miller 
H. Lloyd 

H. Lloyd 

G. A. Miller 
G. A. Miller 
N. Criddle 

G. A. Miller 
N. Criddle 

G. A. Miller* 
N. Criddle 

E. M. Kindle* 


es} 
= 
a 
5 


M 

Ib, 

L 

F 

F 

. M. Sternberg* 
wiePatch= 

. E. Wilson 

. M. Sternberg 
. E. Wilson 

. M. Sternberg 
. E. Wilson 


erie fe) ea19 ke (Viel =" eafevol esli@l 


Qt 
= 
fe) 
= 


P. A. Taverner 
H. Groh 

P. A. Taverner 
P. A. Taverner 
R. DeLury 


Treasurer Secretary(ies)? 
A. Gibson T. E. Clarke 

A. Gibson T. E. Clarke 

A. Gibson E. Clarke 

H. Groh J. J. Carter 

W. T. Macoun D. E. Blackader 
W. T. Macoun D. E. Blackader 
W. T. Macoun E. D. Eddy 


G. LaLacheur 
G. LaLacheur 
J. R. Dymond 
J. R. Dymond* 
F. W. Waugh 

E. B. Crampe* 
C. B. Hutchings 
C. B. Hutchings 
C. B. Hutchings 
B. A. Fauvel 


B. A. Fauvel 


B. A. Fauvel 
B. A. Fauvel 
B. A. Fauvel 


W. Lloyd 


W. Lloyd 
W. Lloyd 
W. Lloyd 
W. Lloyd 
W. Lloyd 
W. Lloyd 


W. Lloyd 


J. F Watson E. D. Eddy 


A. Halkett* 

G. O. McMillan 
L. D. Burling 
L. D. Burling 
C. L. Patch 

C. L. Patch 

C. L. Patch 

C. L. Patch 

C. L. Patch 

C. L. Patch 


J. FE. Wright 


J. E. Wright 


J. F Wright 
J. F. Wright 
J. F Wright 


B. A. Fauvel 


B. A. Fauvel 


G. S. Postethwaite 


G. S. Lewis 


G. S. Lewis 


G. S. Lewis 


P. Whitehurst 


P. Whitehurst 


36 


1937 


1938 


1939 
1940 
1941 
1942 
1943 
1944 
1945 
1946 
1947 
1948 
1949 
1950 
1951 
1952 
1953 
1954 
1955 
1956 
1957 
1958 
1959 
1960 


1961 


1962 
1963 
1964 
1965 
1966 


1967 


THE CANADIAN FIELD-NATURALIST 


President 


H. Groh 


P. A. Taverner 


A. E. Porsild 
A. E. Porsild 
H. G. Crawford 
H. G. Crawford 
D. Leechman 
D. Leechman 
FE. Banim 

F. E. Banim 

W. H. Lanceley 
W. H. Lanceley 
P. Snure 

P. Snure 

J. W. Groves 

J. W. Groves 

R. Frith 

R. Frith 

W. K. W. Baldwin 
W. K. W. Baldwin 
L. S. Russell 

L. S. Russell 

E. L. Bousfield 
E. L. Bousfield 


W. W. Mair* 
D. R. Beckett* 


D. R. Beckett 
D.R. Beckett 
G. McGee 
G. McGee 
G. McGee 


H. Mackenzie 


Vice-President(s)! 


P. A. Taverner 
R. DeLury 

R. DeLury* 

H. G. Crawford* 
A. E. Porsild 

H. G. Crawford 
D. Leechman 

H. G. Crawford 
D. Leechman 

D. Leechman 


D. Leechman 
F. E. Banim 

F. E. Banim 

W. H. Lanceley 
F. E. Banim 

W. H. Lanceley 
W. H. Lanceley 
A. L. Rand 

W. H. Lanceley 
A. L. Rand 

A. L. Rand 

P. Snure 

P. Snure 

J. W. Groves 

J. W. Groves 
R. Frith 

J. W. Groves 
R. Frith 

R. Frith 

W. K. W. Baldwin 


K. W. Baldwin 
Senn 

K. W. Baldwin 
E. Bourguignon 
Russell 
Bousfield 
Russell 
Bousfield 
B 

B 


IL, 

E 

Le 

E ousfield 
leakney 

Bousfield 

Bleakney 

W. W. Mair 

D. R. Beckett 

W. W. Mair 

D. R. Beckett 

D. R. Beckett* 

V. Solman* 

E. Bousfield 

W. W. Mair 

D. A. Smith 

W. W. Mair 

G. McGee 

W. W. Mair 

G. R. Hanes 

W. W. Mair 

F. R. Cook 

F. R. Cook 

H. Mackenzie 

J. C. Woolley 

T. Mosquin 


W. 
H. 
W. 
A. 
9. 
a 
Sy 
Bas 
Hf 
MS 
ae 
ssh 


J 
E 
J 


Treasurer 


W. Lloyd 


W. Lloyd 


W. Lloyd 
W. Lloyd 
W. Lloyd 
C. H. D. Clarke 
I. Conners 
I. Conners 
I. Conners 
I. Conners 
C. Frankton 
C. Frankton 
C. Frankton 
C. Frankton 
R. J. Moore 
R. J. Moore 


R. J. Moore 


R. J. Moore 
R. J. Moore 
R. J. Moore 
R. J. Moore 
R. J. Moore 
J. M. Gillett 
J. M. Gillett 


J. M. Gillett 


A. Banning 
A. Banning 
A. Banning 
A. Banning 
R. D. Wainwright 


L. G. Howden 


Vol. 118 


Secretary(ies)” 
P. Whitehurst* 


C. W. Lounsbury* 
C. W. Lounsbury 
C. W. Lounsbury 
C. W. Lounsbury 
C. W. Lounsbury 
J. W. Groves 

J. W. Groves 

J. W. Groves 

J. W. Groves 

O. H. Hewitt 

O. H. Hewitt 

H. J. Scoggan 
H. J. Scoggan 

J. Scoggan 

J. Scoggan 


J. Scoggan 


al 


. Scoggan 
J. Scoggan 
. Scoggan 
J. Scoggan 
J. Scoggan 
M. Banfield 
M. Banfield 


M. Banfield 


ee ee eee 
a 


M. Banfield 


A. M. Banfield 


D. A. Smith 


A. W. Rathwell 


A. W. Rathwell 


A. W. Rathwell 


A. W. Rathwell 


2004 


1968 


1969 
1970 
1971 
1972 
1973 
1974 
1975 
1976 
1977 
1978 
1979 
1980 
1981 
1982 


1983 


1984 


1985 
1986 
1987 


1988 


1989 
1990 


199] 


1992 
1993 
1994 


1995 


BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 


President 


H. Mackenzie 


H. Mackenzie 
T. Mosquin 
T. Mosquin 
S. Thomson 
I. M. Brodo 
I. M. Brodo 
E. C. D. Todd 


E. C. D. Todd 


R. A. Foxall 


R. A. Foxall 
R. Taylor 

R. Taylor 

R. Taylor 

D. F. Brunton 


D. F. Brunton 


E. F. Pope 


E. F. Pope 
W. Gummer 
W. Gummer 


J. Harrison 


J. Harrison 
J. Harrison 


R. John* 


E. F. Pope 
ESE Pope 
E. EF. Pope 


E. F. Pope 


Vice-President(s)! 


T. Mosquin* 
J. Tener* 

G. McGee* 

T. Mosquin 
W. A. Holland 
W. A. Holland 
S. Thomson 
S. Thomson 

I. M. Brodo 

I. M. Brodo 
E. C. D. Todd 
E. C. D. Todd 


E. C. D. Todd 
R. A. Foxall 
R. A. Foxall 
R. Taylor 

R. Taylor 

C. Gilliatt 

H. L. Dickson 


H. L. Dickson* 
D. F. Brunton* 
C. G. Gruchy 
P. M. Catling 
C. G. Gruchy* 
ESE; Pope* 

P. M. Catling 
W. Gummer 
W. P. Arthurs 


W. Gummer 
W. P. Arthurs* 
B. A. Campbell 
J. Harrison 

D. F. Brunton 
J. Harrison 

D. F. Brunton* 
K. Strang* 

J. Harrison 

R. John 

K. Strang 

R. John 

D. Cuddy 

E. Fox* 

S. Blight* 


vacant 
M. Murphy 


M. Murphy 
D. Moore 
M. Murphy 
D. Moore 


Treasurer 


37 


Secretary(ies)? 


L. G. Howden 


F. M. Brigham 
F. M. Brigham 
F. M. Brigham 
P. Kevin 

C. Gruchy 

C. Gruchy 

P. J. Sims 

P. J. Sims 

B. Henson 

B. Henson 

B. Henson 

B. Henson 

B. Henson 

P. D. M. Ward 


P. D. M. Ward 


P. D. M. Ward 


P. D. M. Ward 
P. D. M. Ward 
F. Valentine 


F. Valentine* 


J. Gehr 
M. Scromeda 


G. Marston 


G. Marston 
G. Marston 
G. Marston 


G. Marston 


A. W. Rathwell 


A. W. Rathwell 
A. W. Rathwell 
A. W. Rathwell 
A. W. Rathwell 


A. J. Erskine (RS) 
A. H. Reddoch (CS) 
A. J. Erskine (RS) 
J. D. Lafontaine (CS) 
A. J. Erskine (RS) 
C. Gruchy (CS) 

P. J. Narraway (CS) 
A. J. Erskine (RS) 
D. Laubitz (RS) 

S. Armstrong (CS) 
D. Laubitz (RS) 

S. Armstrong (CS) 
D. Laubitz (RS) 

V. Hume (CS) 

D. F. Brunton (RS) 
E. F. Pope (CS) 

E. F. Pope (RS) 

W. Gummer (CS) 
E. F. Pope (RS) 

W. Gummer (CS) 
E. F. Pope (RS) 

W. Gummer (CS) 


G. Hamre (RS)* 

A. Martell (CS)* 

B. Martin (RS & CS)* 
A. Martell (CS)* 

B. A. Campbell (CS)* 
B. J. Martin (CS) 

E. Bottomley (RS) 
B. A. Campbell (CS) 
M. Coleman (RS) 

D. F. Brunton (RS)* 
R. John (RS)* 

B. A. Campbell (CS) 
D. Duchesne (RS) 
M. Aksim (CS) 

E. Fox (RS) 

E. Evans (CS) 

E. Evans (CS) 

D. Furlong (RS)* 

C. Firth (RS)* 

N. Stow (RS)* 

E. Fox (RS)* 

E. Evans (CS) 

C. Clark (RS) 

S. Gawn (RS) 

E. Evans (CS) 

S. Gawn (RS) 

E. Evans (CS) 

D. Smythe (RS) 

E. Evans (CS) 


38 THE CANADIAN FIELD-NATURALIST Vol. 118 
President Vice-President(s)! Treasurer Secretary(ies)” 
1996 D. Moore M. Murphy G. Marston D. Smythe (RS) 
E. Evans (CS) 
1997 D. Moore M. Murphy S. Shaw* D. Smythe (RS) 
T. Reeve* L. Cairnie (CS) 
1998 D. Moore E. Zurbrigg S. Shaw G. McNulty (RS) 
D. Smythe vacant (CS)* 
1999 D. Moore E. Zurbrigg ESE: Pope? G. McNulty (RS) 
D. Smythe vacant (CS)* 
2000 E. Zurbrigg R. John E. F. Pope J. Martens 
2001 E. Zurbrigg R. John E. F. Pope K. Allison 
2002 E. Zurbrigg R. John E. F. Pope K. Allison 
2003 G. McNulty M. R. Murphy E. F. Pope S. L. Bourque 
G. Marston 
2004 M. R. Murphy G. Marston E. F. Pope S. L. Bourque 


* served only a portion of this term; RS — Recording Secretary CS — Corresponding Secretary 


' First and Second Vice-President positions existed in most but not all terms. 

2 Secretary and Treasurer positions were combined until 1882; 

> The OFNC “year” was rearranged in 1919 to coincide with the calendar year rather than the fiscal year, resulting in a 
1918-1919 year ending in March 1919, a9 month “year” for the remainder of 1919, and normal calendar years thereafter. 

* Corresponding Secretary position left vacant for two years then eliminated; Recording Secretary position continues. 


Continuing Environmental Change — An Example from Nova Scotia 


EDMUND S. TELFER 


Scientist Emeritus, Canadian Wildlife Service, Environment Canada 
Current address: 3582-42 Street, Edmonton, Alberta, T6L 5A1 Canada 


Telfer, Edmund S. 2004. Continuing environmental change — an example from Nova Scotia. Canadian Field-Naturalist 
118(1): 39-44. 


Information from personal experience, from community elders and published literature served as a basis for evaluating envi- 
ronmental changes in the District of North Queens and adjacent areas of Southwestern Nova Scotia over the past century. Major 
events included disappearance of the Caribou (Rangifer tarandus), the arrival of White-tailed Deer (Odocoileus virginianus), 
the severe reduction of Canada Yew (Taxus canadensis), disappearance of Lynx (Lynx canadensis), a major dieoff of Striped 
Skunks (Mephitis mephitis), decline of American Beech (Fagus grandifolia), the loss of mature birch (Betula spp.), the severe 
reduction of Moose (Alces alces), the arrival of the American Dog Tick (Dermacentor variabilis) and Coyotes (Canis latrans), 
and the restoration of Beaver (Castor canadensis). The proximate cause of many of those changes were plant and animal 
disease, while the ultimate causes were naturally occurring animal range expansion and human impacts. The warming of the 
climate over the past 150 years probably played a role. The nature and timing of the events could not have been predicted. 


Key Words: Nova Scotia, plant and animal diseases, change in biota, climate change, adaptive management. 


Here, I present and discuss an example of long-term 
environmental change. It is a narrative account of 
changes in the occurrence of some mammals and plants 
in my home area, the District of North Queens, and 
surrounding portions of the interior of southwestern 
Nova Scotia. I personally observed these changes or 
heard about them from my father and other community 
elders and further document them by reference to the 
available literature. I discuss the changes in the light 
of current resource management philosophy. Since my 
father was born in 1882 and was a teenager in the 1890s, 
the changes noted cover a century. 

The District of North Queens lies in the interior of the 
southwestern peninsula of Nova Scotia and is centred at 
approximately 65°09'W and 44°18'N. It is an area of 
scattered farms and homesteads where the population 
has subsisted for 200 years on a mix of agriculture and 
forestry. The district contains the western portion of a 
swarm of glacially-formed drumlin hills that occur in 
Lunenburg and Queens counties (Roland 1982). Drum- 
lins provide arable land but comprise a small propor- 
tion of the land area. Most of the district is rocky, for- 
ested terrain that, even where cultivatable and produc- 
tive, was difficult for early settlers to clear. The remain- 
der of the interior of western Nova Scotia is a complex 
of water bodies, bogs, fens, barrens and stony soils 
supporting Acadian forests of mixed conifers and decid- 
uous tree species (Rowe 1972). When my father was 
born the area contained considerably more crop agri- 
culture and improved pasture than today. For the County 
of Queens as a whole, approximately 30 000 acres 
(12 140 ha) were under agriculture in 1870 (Canada 
1871). This area dropped to 7410 acres (2999 ha) by 
1966 (Nova Scotia 1966). However, those were small 
proportions of the total area, 4.4% in 1870 and 1.1% 


in 1966. Because the terrain is very complex, and the 
forest composed of many small stands, with the whole 
divided into small private land parcels, timber harvest 
has consisted of cutting small patches or individual trees 
as a market or need for the wood occurred. However, in 
recent years there have been larger clearcuts and some 
conversion of mixed forest to conifer plantations on 
industrial ownerships. The decline in the area of mixed- 
wood has been counterbalanced to an unknown extent 
by reversion of agricultural land to mixedwood forest 
and by improved fire protection (Telfer 1971). 


Mammals 

In the 1880s Caribou Rangifer tarandus, were still 
common in unsettled parts of Nova Scotia (Rand 1933; 
Sheldon 1936). An old hunter (the late F. M. Forrest, 
personal communication) told me of a trip he made 
when he was 16 years old (that would have been in 
1888) to find Caribou. Somewhere at the west side of 
the present day Kejimkujik National Park he found a 
herd on a large barren and had a good chance to ob- 
serve them and to shoot a fat yearling to replenish his 
provisions. Within 20 years Caribou appear to have been 
gone from North Queens. Another old woodsman (the 
late E. B. Smith, Sr., personal communication) told me 
that when he began to guide sportsmen in the early 
1900s there were still broad trails beaten though the 
woods in all directions in the remote parts of the District 
that had been made and kept open by Caribou. Sheldon 
(1936) also commented on the Caribou trails. By my 
time the Caribou were long gone and their trails had 
faded away before the encroaching forest. Caribou 
suffered from logging and burning on their winter 
ranges and from poorly controlled hunting. However, 
the probable final cause of Caribou extirpation was 


39 


40 THE CANADIAN FIELD-NATURALIST 


meningeal worm, Parelaphostrongylus tenuis, intro- 
duced by White-tailed Deer, Odocoileus virginianus 
(Benson and Dodds 1977). 

The loss of a large mammal like Caribou from the 
regional fauna quite probably had at least minor im- 
pacts on other plants and animals living in the area. I 
have not been able to find any records of any obser- 
vations of such impacts. However, Caribou move fast 
and far and feed while moving so they spread their 
use of vegetation out in time and space (Miller 1982). 
The relatively small population of Caribou in Nova 
Scotia would have had slight impact on other species. 

Moose, Alces alces, were distributed throughout 
Nova Scotia and were the common game animal in 
my father’s youth. However, after 1926 they were 
observed to be dying (Sheldon 1936; Benson 1958). 
The sick Moose exhibited symptoms similar to those 
later described by Anderson (1965) as resulting from 
infestation by the meningeal worm (Parelaphostron- 
gylus tenuis). This organism was later found in Nova 
Scotia Moose by Smith et. al (1964). Parelaphostron- 
gylus tenuis is relatively common in White-tailed Deer. 
It does little harm to whitetails but may cause heavy 
mortality among other species of the deer family 
(Thomas and Dodds 1988). By the 1940s, Moose were 
dying in large numbers from the condition which had 
come to be called “Moose Sickness” (Benson and 
Dodds 1977). The animals showed evidence of paral- 
ysis, blindness and other neurological symptoms al- 
though they were often in good physical condition 
otherwise. Annual reports of the Nova Scotia Depart- 
ment of Lands and Forests commented on the dead 
Moose. The 1949 report was typical, “Nearly all 
Rangers report moose as being in a healthier condi- 
tion which is perhaps indicated by the decrease in the 
number of moose which were found dead. This number 
was 75, a decrease of 74 from last year” (Nova Scotia 
1949: 65). Of course only a fraction of the dead Moose 
would have come to the attention of the authorities. 
Moose almost vanished from southwestern Nova Scotia 
by the 1960s but recovered slightly following later 
declines in deer numbers. 

Although P. tenuis was found in some Nova Scotia 
Moose in the 1960s and has been shown to cause a 
“moose sickness” type of illness, that does not prove 
that all the animals that perished in the dieoff of 
Nova Scotia Moose in the 1930s and 1940s died of 
meningeal worm infection. However, the similarity of 
the symptoms displayed by the sick Moose suggests 
that most died of the same cause. Moose have been 
found to suffer from neurological symptoms where no 
P. tenuis has been observed in autopsies (Tony Nette, 
personal communication). Those symptoms have been 
recently linked to deficiencies of cobalt and vitamin 
B,, (Frank et al. 2004). However, it seems unlikely 
that a nutritional deficiency would cause a catastrophic 
dieoff like that reported in Moose in Nova Scotia in 
the 1940s, and the coincidence of the dieoff with the 


Vol. 118 


post-introduction boom in White-tailed Deer num- 
bers suggests that P. tenuis was the principal cause. 

My father was an adult before deer showed up in 
North Queens. They had been spreading across Nova 
Scotia with the help of some introductions, and by the 
late 1800s their numbers were increasing (Rand 1933). 
There were many reasons for the increase. Possibly 
the most decisive was the climatic warming that set in 
about 1850 (Hawboldt 1952; Lamb 1982). The first 
deer hunting season in Nova Scotia was in 1916 and by 
the 1930s deer were common (Sheldon 1936; Benson 
and Dodds 1977). Over 40 000 were reported killed in 
Nova Scotia in one year in the early 1950s — more than 
two for every square mile in the province (Benson and 
Dodds 1977). Deer remained numerous until the unusu- 
ally severe winter of 1955-1956 caused a major dieoff 
of deer in western Nova Scotia. Since then they have 
fluctuated with winter conditions. 

Beavers, Castor canadensis, were not mentioned by 
my father as a species that he encountered in North 
Queens as a young man in the 1890s and the early 
1900s. The animals had been trapped almost to extirpa- 
tion in Nova Scotia during the preceding century. How- 
ever, in 1907 Beavers were given complete protection 
(Wood 1973) and the Department of Lands and For- 
ests staff began live-trapping and transplanting them 
from the few remaining populations to unoccupied 
parts of the province. 

One cold autumn dusk when I was about five years 
old my father took me on a short walk to a small lake 
where Beavers had been released. We saw a new dam 
blocking the outlet stream and a new house with 
fresh mud on it. The Beavers had made themselves at 
home. They soon became abundant in North Queens 
and remained so until after 1945. Limited trapping was 
allowed after that time, reducing Beaver numbers and 
the dense population of animals had reduced their food 
supply along streams and lakes. However, Beavers have 
continued to occur in substantial numbers in North 
Queens (Wood 1973). 

Hunting with hounds was a favourite activity of my 
father and his friends. They hunted Red Foxes (Vulpes 
vulpes) and what they referred to as “wildcats”. Most 
of the cats were Bobcats (Lynx rufus) but my father 
also described killing cats that were over six feet long 
from rear toes to front toes when held up by their 
hind legs. Those large, rangy animals were probably 
Lynx. More (1873) stated that there were “lucifees” 
(Loup cervier, Lynx) and “little lucifees” in Queens 
County. However, the Lynx had become rare or extir- 
pated by the 1930s (Rand 1933; Sheldon 1936) and 
is now absent from the region (McCord and Cardoza 
1982). 

According to local tradition, there were no Striped 
Skunks, Mephitis mephitis, in North Queens when the 
area was first settled. The species is not mentioned by 
More (1873) who, as a land surveyor, was very famil- 
iar with the mammals of the region. It was said that 


2004 


sometime in the mid-1800s someone brought a dead, 
frozen animal to a local blacksmith shop. None of the 
farmers who happened to be there could identify it. 
However, a few days later someone passed who had 
worked in the United States and who identified it as a 
skunk. By my father’s time skunks were common. 
However, by the 1930s, skunks were gone, probably 
due to an epidemic of distemper (Dodds 1969) but 
the decline in agricultural land use may also have had 
a negative impact on this animal of farms and forest 
edge. They were found in Nova Scotia only along the 
New Brunswick border but by the 1960s they had 
once again begun to spread into central areas of the 
province (Dodds 1969). None were present in Kejim- 
kujik National Park in the early 1970s (Wood 1973). 
However, in 2001 I observed a skunk near Ten Mile 
Lake, in Queens County. 

My father would have had difficulty in believing that 
the howl of the Coyote (Canis latrans) could ever be 
heard in North Queens. Yet in the 1970s Coyotes ar- 
rived in Nova Scotia. In North Queens, the popula- 
tion of White-tailed Deer in Kejimkujik National Park 
was apparently reduced by Coyotes which were com- 
monly seen along Park roads and heard on still nights 
throughout the area (Patterson 1994). The impact of 
Coyotes not only on deer but on competing Red Foxes, 
on Snowshoe Hares (Lepus americanus), microtine 
rodents, grouse (Bonasa umbellus and Canachites can- 
adensis) and other ground-nesting birds must have 
been considerable. 


Plants 

Before the arrival of White-tailed Deer, the forest 
floor in mature spruce-hemlock stands was carpeted 
with Canada Yew, Taxus canadensis. This evergreen 
shrub is highly palatable to deer and by the 1940s it 
had been eliminated from the forest. In the 1960s one 
experienced biologist (the late Harrison F. Lewis, 
personal communication) told me that the only sur- 
viving yew bush that he knew of was growing beside 
the outhouse at a warden cabin where one branch had 
grown through the wall of the well-ventilated facility 
and had thus escaped browsing. 

One of the most important trees of North Queens is 
the White Pine (Pinus strobus) (Rowe 1972). White 
Pines were a major resource in North Queens histori- 
cally (More 1873). However, in my father’s time re- 
generating White Pine was increasingly attacked by 
the White Pine Weevil (Pissodes strobi). That insect 
kills the leaders on the top of the tree, creating crooked 
stems or, at worst, “cabbage trees” whose lower branch- 
es continually grow up to take the place of the weevil- 
killed central stem. White Pine Weevil is a native North 
American insect. However, the first official records of 
its presence in the Maritimes date from about 1920 
(Martineau 1984). The insect quickly spread throughout 
the range of the White Pine including North Queens. 
The spread of the weevil was facilitated by the exten- 


TELFER: CONTINUING ENVIRONMENTAL CHANGE — AN EXAMPLE 4| 


sive areas of regenerating forest created by logging, fire 
and the invasion of old fields by White Pine (Belyea 
and Sullivan 1956). By my time the value of the White 
Pine resource in North Queens was greatly reduced 
and crooked, multi-stemmed pine occupied much space 
in the forest. White Pine remains an important tree in 
the district but the weevil attacks have reduced the 
value of many stems for lumber. 

The American Beech, Fagus grandifolia, was com- 
mon in North Queens when my father was a boy. Pure 
beech stands occupied the ridge tops and beech trees 
were also scattered throughout the hardwood and 
mixedwood stands. Beech nuts were always an impor- 
tant seasonal food source for wildlife. Black Bears 
(Ursus americanus), especially, sought them out for 
a high fat and protein food before hibernation. When 
whitetails arrived in Nova Scotia they quickly became 
the most important consumers of the dwindling supply 
of nuts. Loucks (1962) cited evidence that beech was 
formerly more common in southwestern Nova Scotia. 
He believed that wildfire had reduced the occurrence 
of the species. Early in the 1900s the Woolly Beech 
Scale (Cryptococcus spp.), an insect that spreads Beech 
Nectria Fungus (Nectria coecinea var. faginata), ar- 
rived in Nova Scotia, probably an accidental intro- 
duction from Europe (Boyce 1961). The fungus gains 
entry to the tree through holes made by the Woolly 
Beech Scale. Saunders (1970) stated that Woolly 
Beech Scale entered Nova Scoria from Europe about 
1890 and that the fungus also entered North America 
through Nova Scotia in about 1929. Nectria disease 
debilitates beech by invading the bark and cambium, 
eventually killing the tree. Roland and Smith (1969: 
page 341) noted that “All the beech in N.S. is severe- 
ly affected by the Nectria beech canker.” Most beech 
in North Queens was unthrifty and dying by the mid- 
twentieth century and in all cases moribund stems were 
thoroughly infested with Nectria cankers. Beech have 
declined substantially (Telfer 1971). They have been 
replaced by species like Sugar Maple (Acer saccha- 
rum) and Red Maple (Acer rubrum), Hemlock (Tsuga 
canadensis) and Red Spruce (Picea rubens). 

White Birch (Betula papyrifera) and Yellow Birch 
(Betula alleghaniensis) were prominent trees in the 
North Queens forest when my father was a boy. In- 
deed the old farmhouse in which he was born and died 
was sheathed with bark of the white birch between the 
wallboards and the shingles. In the mid-1930s a die- 
back was observed in the birch (Hawboldt 1952. 
Mature stands all died. This phenomenon occurred all 
over northeastern North America. 

Temperatures were high in the 1930s following a 
steady rise from Little Ice Age conditions that existed 
before the 1850s (Hawboldt 1952). One hypothetical 
cause of birch mortality was that the warming climate 
altered the species composition of the mycorrhizal 
flora in the soil to the detriment of species that could 
enter into symbiotic relationships with birch roots 


42 THE CANADIAN FIELD-NATURALIST 


(Hawboldt 1952). Birch would thus have suffered from 
nutrient deficiencies. Although birch regeneration sur- 
vived, live mature trees were very scarce by my time. 
One would have had to travel a long way to collect 
enough bark to sheath a house. In the 1960s I worked 
on forest inventories in a remote part of the District 
where the mature forest had formerly been dominated 
by a mixture of birches, Red Spruce and Hemlock 
(Fernow 1912), but by then the birches were rotten 
snags and Red Spruce, White Pine, Balsam Fir (Abies 
balsamifera) and Red Maple were sprouting in the 
gaps created by their death. 


Arachnids 

An invading organism that caused excitement in 
North Queens in the mid-twentieth century was the 
American Dog Tick (Dermacentor variabilis). Opinion 
is divided on whether the ticks were relict in a small 
area in Western Nova Scotia or if they were introduced. 
One woodsman told me of encountering ticks along a 
small section of the Tusket River, near the western 
end of Nova Scotia, during a canoe trip early in the 
1900s (the late F. M. Forrest, personal communi- 
cation). The distribution pattern shown by Dodds et 
al. (1969) suggests dispersion from an epicentre of 
infection. The Tusket River flows through the area of 
earliest distribution. By the 1930s loggers working in 
the western part of North Queens began to complain 
of tick bites. Ticks advanced on a broad front across 
the district in the late 1940s and the 1950s, causing 
consternation among the people who were unused to 
them. Since the 1960s, American Dog Ticks have con- 
tinued to spread eastward in Nova Scotia (Andrew 
Hebda, personal communication). 


Weather 

Nova Scotia is on the track of the equatorial Atlantic 
hurricanes. North Queens has been occasionally lashed 
by at least the fringes of those storms. In 1939, the area 
was hit by a hurricane that blew down many trees but 
did limited damage in closed stands. However, in 1954, 
a powerful hurricane, code named “Edna”, struck North 
Queens (Hawboldt and Bulmer 1958). Mature and old 
forests suffered particularly. Hurricane Edna was fol- 
lowed by a campaign of salvage logging as landown- 
ers tried to recover as much of the blowdown as pos- 
sible. The forest openings were invaded by regeneration 
of Red Spruce, Red Maple, White Pine and particu- 
larly by Balsam Fir in place of the dominant Hemlock 
of the former stands. 


Prediction and Management 

Causes of some of the observed changes were nat- 
ural, like birch dieback and distemper, while others 
were due at least in part to human intervention. There 
were some introductions of White-tailed Deer to Nova 
Scotia that probably accelerated their spread. Logging, 
small farming and attendant wildfires no doubt in- 


Vol. 118 


creased the amount of suitable habitat for deer and also 
the area occupied by White Pine regeneration, the 
habitat of the White Pine Weevil. Hunting probably 
hastened the extirpation of Caribou and Lynx and the 
near extirpation of beaver. Coyotes may have been 
able to invade eastern North America partly because 
European settlers extirpated Grey Wolves, Canis lupus, 
from regions to the west of Nova Scotia. Ticks and 
the insect vector and the fungus involved in the beech 
Nectria disease may have been accidentally introduced. 
Climatic warming over the past 150 years interacted in 
a complex way with other causes of ecological change. 

The most important factors causing change to the 
terrestrial biota in North Queens operated haphazardly. 
Most were outbreaks of plant and animal diseases 
and parasites. Unpredictable outbreaks of disease are 
sometimes not considered to be very important by re- 
newable resource managers simply because they are 
random and basically unforeseeable. Also, the educa- 
tion of resource managers usually focuses on other 
topics. However, Hurricane Edna represented another 
kind of random event. There was sufficient evidence 
of the occasional occurrence of such severe hurricanes 
that shrewd ecosystem managers would probably have 
allowed for their possible occurrence when preparing 
management plans. 

The observed ecological changes in North Queens 
did not damage basic ecological functions like produc- 
tion, herbivory and predation. However, the suite of 
species involved changed. White-tailed Deer to some 
extent replaced Moose and Caribou as consumers 
while Coyotes became important predators as Lynx 
had been previously. After decades of absence, Beavers 
were brought back in numbers by deliberate manage- 
ment. Energy flow through the ecosystem was unim- 
paired but flowed through somewhat different chan- 
nels. The observed changes exemplify the fluidity and 
unpredictability of nature (Botkin 1990). 

What might have happened if, beginning in the 
1890s, the forests and wildlife of North Queens had 
been under a modern regime of ecosystem manage- 
ment? Foresters might have made management plans 
aiming at production of a steady supply of birch logs. 
All would have been well for the first fifty years, then 
they would have found that the birch growing stock 
was dying and that spruce and fir were taking its place. 
As time went on the beech would also have been lost 
as a commercial species. Forest management would 
have had to be altered, as it indeed was following Hur- 
ricane Edna when managers were forced to revise their 
thinking about harvesting techniques (Johnson 1986) 
and deal with storm-caused openings and changes in 
forest composition. Conversely, the patchy nature of the 
storm-damaged forests provided improved habitat for 
White-tailed Deer and Moose. 

Wildlife managers would have found a major ungu- 
late species, the Caribou, suddenly declining. Even if 
the meningeal worm problem had been known, would 


<2 


2004 


managers have tried solve it by exterminating the 
White-tailed Deer population? Whitetails were by then 
spread thinly over the entire province. By the time 
Moose also declined there were so many deer that erad- 
ication would have been impossible. Even without par- 
asites, how would biodiversity managers have viewed 
the invasion of exotic whitetails especially since they 
were also the culprits in the extirpation of the Canada 
Yew? Managers would have been largely helpless to 
protect the yew except possibly by fencing small res- 
erved patches. Although “ecosystem” preservation is 
often a suitable management approach to maintain 
threatened species, it would have been counterproduc- 
tive in this case because old hemlock/spruce forest is 
preferred winter habitat for whitetails as well as the 
site of best yew growth (Schierbeck 1931). Lynx still 
remain on Cape Breton Island so protection from hunt- 
ing might have been adequate to recover the species 
on the mainland of Nova Scotia as it did for the 
Beaver. Because skunks appear to have died from dis- 
temper, reintroduction of healthy animals following 
the dieoff might also have re-established viable popu- 
lations and prevented local extirpation. 

Our hypothetical ecosystem managers would have 
emerged from the hundred-year period somewhat bat- 
tered. They would have lost a member of the charis- 
matic megafauna, the Caribou. They would no doubt 
have been chagrined by the fact that the district had 
been overrun by the exotic White-tailed Deer and Coy- 
otes, not to mention the obnoxious dog ticks. Some 
major tree species would also have been drastically 
reduced. However, a major understorey shrub and two 
mammal species might have been rescued and one 
mammal species, the Beaver, was indeed returned to 
the area by management. Would the public consider 
that money spent trying to maintain the ecosystems 
as they were in the 1890s to have been well spent as 
part of a worthwhile effort to “preserve” “nature” in 
the region? Or would managers simply be seen as 
having cast themselves in front of the juggernaut of 
inevitable ecological change? 

In the 1890s the coming changes and their impact 
could not have been foreseen. If managers at that time 
had set goals like doubling the sustainable harvest of 
White Birch and developing an industry to use it, or 
if they had set out to double the Caribou population to 
satisfy outfitters, they would have been in for a rough 
time. Yet, such factory-type production goals are often 
set in resource management. 

This example highlights the significance of scale 
in biodiversity conservation. Would it have been worth- 
while to try to keep species that are still common else- 
where in North America, like Caribou, Lynx, Striped 
Skunks, and Canada Yew, in the district or even in Nova 
Scotia? At what price should we try to keep everything 
that was present historically in a local or regional biota? 
Or should the waxing and waning of species’ ranges 
be accepted as inevitable? 


TELFER: CONTINUING ENVIRONMENTAL CHANGE — AN EXAMPLE 43 


Although such a comprehensive management ap- 
proach would have been unthinkable in the 1890s, a 
program of adaptive management (Walters 1986) would 
have been appropriate. With the adaptive model in 
mind, managers might have been aware of the likeli- 
hood of natural variability and therefore cautious in 
goal setting. They could have initiated a program of 
monitoring that might have picked up changes in spe- 
cies status in their early stages when some prediction 
of the magnitude of upcoming impacts could have been 
made. While little could have been done to change the 
course of events in the short term, a more comprehen- 
sive and quantitative understanding of the operation of 
nature might have led to effective interventions later 
on. 

The haphazard nature of the changes in the ecology 
of North Queens during the past hundred years under- 
lines the difficulty of predicting what will happen next 
in nature. One is reminded of the soldier’s axiom re- 
garding shellfire, “You won’t hear the one that gets 
you”. The example of North Queens demonstrates the 
inevitability of environmental change and emphasizes 
the value of a flexible approach to forest and wildlife 
management. 


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Odocoileus virginianus, by Coyotes, Canis latrans, in 
Nova Scotia. Canadian Field-Naturalist 108: 484-487. 


Vol. 118 


Rand, A. L. 1933. Notes on the mammals of the interior of 
western Nova Scotia. Canadian Field-Naturalist 47: 41- 
60. 

Roland, A. E. 1982. Geological background and physio- 
graphy of Nova Scotia. Nova Scotia Institute of Science, 
Halifax. 311 pages. 

Roland, A. E., and E. C. Smith. 1969. The flora of Nova 
Scotia. The Nova Scotia Museum. 743 pages. 

Rowe, J. S. 1972. Forest regions of Canada. Department of 
the Environment, Canadian Forest Service Publication 
1300. 172 pages plus map. 

Saunders, G. L. 1970. Trees of Nova Scotia. Bulletin 37, 
Nova Scotia Department of Lands and Forests. 102 pages. 

Schierbeck, O. 1931. Forestry vs. game cover. Canadian 
Field-Naturalist 45: 28-30. 

Sheldon, C. 1936. The mammals of Lake Kedgemakooge and 
vicinity, Nova Scotia. Journal of Mammalogy 17: 207-215. 

Smith, H. J., R. McG. Archibald, and A. H. Corner. 1964. 
Elaphostrongylosis in Maritime moose and deer. Canadian 
Veterinary Journal 5: 287-296. 

Telfer, E. S. 1971. Changes in carrying capacity of deer range 
in western Nova Scotia. Canadian Field-Naturalist 85: 231- 
234. 

Thomas, J. E., and D. G. Dodds. 1988. Brainworm, Pare- 
laphostrongylus tenuis, in moose, Alces alces, and white- 
tailed deer, Odocoileus virginianus, of Nova Scotia. Cana- 
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Walters, C. 1986. Adaptive management of renewable res- 
ources. Macmillan Publishing Company, New York. 

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ence 27: 43-58 (1971-1973). 


Received 15 February 2002 
Accepted 24 June 2004 


Habitat Segregation Among Songbirds in Old-Growth Boreal 
Mixedwood Forest 


ENID E. CUMMING 
1542 Empress Ave., Saskatoon, Saskatchewan S7K 3G3 Canada 


Cumming, Enid E. 2004. Habitat segregation among songbirds in old-growth boreal mixedwood forest. Canadian Field- 
Naturalist 118(1): 45-55. 


The foraging behaviour of ten species of insectivorous songbirds — Boreal Chickadee (Poecille hudsonicus), Golden-crowned 
Kinglet (Regulus satrapa), Ruby-crowned Kinglet (R. calendula), Blue-headed Vireo (Vireo solitarius), Tennessee (Vermivora 
peregrina), Blackburnian (Dendroica fusca), Magnolia (D. magnolia), Yellow-rumped [Myrtle] (D. coronata), Black-throated 
Green (D. virens), and Bay-breasted (D. castanea) warblers — was observed in the boreal mixedwood forest of Prince Albert 
National Park in central Saskatchewan. Birds segregated their habitat use by preferentially foraging in different tree species, 
and through preferential use of different foraging locations (height and position) within trees. 

White Spruce (Picea glauca) was used more than expected by Yellow-rumped Warblers, and Golden-crowned and Ruby-crowned 
kinglets. Tennessee and Magnolia warblers used White Birch (Betula papyrifera), more than expected and Boreal Chickadees 
and Blue-headed Vireos used Balsam Fir (Abies balsamea) more than expected. Boreal Chickadees, Ruby-crowned Kinglets, 
and Tennessee, Blackburnian and Yellow-rumped warblers all used the bottom part of trees less than expected, while Blue- 
headed Vireos foraged near the top of trees less than expected. Large inner branches were avoided by Tennessee, Blackburnian 
and Yellow-rumped warblers, while Bay-breasted Warblers and Blue-headed Vireos avoided small outer twigs. In conifers, 
Blackburnian Warblers foraged significantly higher in the trees than all other species except Black-throated Green and Bay- 
breasted warblers. Blackburnian Warblers also foraged significantly higher than Blue-headed Vireos and Magnolia Warblers 


in deciduous trees. 


Key Words: Songbirds, behaviour, foraging, boreal forest, old-growth, Prince Albert National Park, Saskatchewan. 


Behavioural interactions and niche partitioning in 
songbirds have received much attention in the literature 
(MacArthur 1958; Sabo 1980; Maurer and Whitmore 
1981). For many boreal songbirds, however, much on 
their basic biology remains poorly understood (Sodhi 
and Paszkowski 1995). Many of the detailed behavioral 
descriptions have come from studies conducted in the 
New England States (Rabenold 1978; Robinson and 
Holmes 1982; Hunt and Flaspohler 1998). This is like- 
ly an incomplete description for some birds, as several 
of the species discussed have their breeding ranges 
almost entirely restricted to the Canadian boreal forest 
(Ficken et al. 1996; Rimmer and McFarland 1998). 
Studies on the southern fringes of the boreal forest, in 
Maine or Minnesota, are not adequate on their own to 
understand boreal ecology as many of the plant and ani- 
mal species are not the same as further north (Rowe 
1972; Acton et al. 1998). Even for wide-ranging species, 
there is evidence that birds do not behave the same 
way and do not use the same kind of habitat in all parts 
of their range (Noon et al. 1980; Collins 1983; Robi- 
chaud and Villard 1999). 

Recent studies in the boreal forest have focused on 
habitat associations, and habitat requirements of song- 
birds (Schieck et al. 1995; Kirk et al. 1996). This infor- 
mation is urgently needed in light of increased pressure 
from large-scale forestry in this region (Cummings et 
al. 1994). There is mounting evidence to show that 
mature and old boreal mixedwood forest support an 
entirely different suite of species not found in younger 


age classes (Stelfox 1995; Cumming and Diamond 
2002). These older age classes of forest are currently 
under a harvesting intensity that is disproportionate to 
their abundance in the landscape (Cummings et al. 
1994; Weyerhaeuser 1998*). 

The goal of this project was to investigate details of 
within-stand habitat segregation among boreal song- 
birds, focusing on the foliage-gleaning songbird guild 
of old (>120 years) boreal mixedwood forest. This type 
and age of forest was chosen as it was biologically the 
most diverse and structurally the most heterogeneous 
habitat in the region (Erskine 1977; Johnson et al. 1995: 
Robichaud and Villard 1999). In addition, mature to old 
(80+ years), mixedwood forest will become increas- 
ingly rare at the landscape level if current land use 
practices (i.e., logging, oil and gas exploration and agri- 
culture) continue at their present rate (Cummings et 
al. 1994: Hobson et al. 2002; Schneider et al. 2002*). 


Study Area 

This study was carried out in the boreal, mixedwood 
forest of Saskatchewan (Kabzems et al. 1986; Acton 
et al. 1998), in Prince Albert National Park (53°35'N, 
106°00'W) during 1990 and 1991 (see Bayne and 
Hobson 1997 for description). Three sites were chosen 
in forest stands that were >40 ha in area and >120 years 
old. Sites were as similar in topography and vegetative 
cover as field conditions allowed. Stands were aged 
by forest inventory maps, and by taking core samples 
from six of the largest trees on each site. Each study 


45 


46 THE CANADIAN FIELD-NATURALIST 


site was surrounded by continuous forest of similar 
type, but younger age. None of the study sites had been 
subjected to forest harvesting, and wildfire was the 
driving force behind forest succession in this area 
(Weir et al. 2000). 

The study sites were structurally very heterogeneous, 
with many tree-fall gaps and many standing dead trees. 
These canopy gaps allowed for regeneration, which 
resulted in a wide range of tree sizes and ages. Domi- 
nant tree species were White Spruce, Picea glauca, 
Balsam Fir, Abies balsamea, and Trembling Aspen, 
Populus tremuloides, with scattered White Birch, Betula 
papyrifera. Average canopy height was 21 metres with 
a maximum height of 38 metres (Cumming 1995). 
White Spruce was the only species that reached heights 
of over 30 metres, with some specimens also having 
diameters in excess of one metre. Understory trees (<12 
metres), were mainly Balsam Fir, while the main 
shrubby species were Balsam Fir, White Birch sap- 
lings, and Alder, A/nus sp. Shrub distribution was 
patchy, with dense patches occurring in tree-fall gaps, 
and scattered shrubs in the rest of the study area. The 
most abundant ground cover species were various 
mosses, Bunchberry (Cornus canadensis), Wild Sarsa- 
sparilla (Aralia nudicaulis), Ostrich Fern (Matteuccia 
struthiopteris) and several horse-tail species (Equi- 
setum Spp.). 


Methods 

Observations on bird behaviour took place from 
late May to early August in 1990 and 1991, and birds 
were observed between 04:00 (dawn) and 21:00 
(dusk). Upon locating a focal bird, continuous obser- 
vations were made using binoculars until the bird 
was lost from sight (Altman 1974; Martin and Batson 
1986). To avoid possible biases, observations that last- 
ed less than five seconds or longer than two minutes 
were not used in the analysis. Bird species, behaviour, 
tree species and bird position in the tree were record- 
ed. Bird height was estimated to within three metres. 
Linear regression analysis (Zar 1996) was used to 
ensure that estimates accurately reflected clinometer- 
measured heights (Cumming 1995). 

In addition to behavioural observation, unlimited 
distance point counts (Bibby et al. 1982) were con- 
ducted twice during the breeding season (June), in 
order to estimate relative abundances of the various 
bird species. The average number of individuals per 
point count for each bird species was obtained by taking 
the maximum number recorded for that species and 
dividing by the number of point counts. Data from the 
three sites were pooled and used as a relative index of 
abundance for the study as a whole. For more details 
on bird species abundances see Cumming and Diamond 
(2002). 

Following the method of MacArthur (1958), con- 
ifers were divided into 15 zones; 5 vertical and 3 hori- 
zontal, to determine in which part of the tree birds were 
foraging (Figure 1). Vertical zones divided the tree into 


Vol. 118 


fifths, with zone | at the top and zone 5 at the bottom. 
Horizontal zones corresponded to the outer part of the 
branch with new growth, fresh needles and small twigs 
(zone T), the middle layer of older needles and larger 
branches (zone M), and the inner part of the branch, 
largely devoid of needles with large diameter branches 
(zone B). A similar system was used for deciduous 
trees except they were divided into three vertical zones 
due to their more globular shape (Figure 2). 

Although many birds exhibit male-female foraging 
differences (Holmes 1986), data for both sexes were 
pooled since the focus of the study was on interspecific 
differences in habitat use and because some species 
in the study were not sexually dimorphic. Bird foraging 
behaviour (tree zone use), did not differ significantly 
between the two species of conifer or between the two 
species of deciduous trees. Therefore, data for each 
bird species was pooled based on either coniferous or 
deciduous trees (Cumming 1995). 

Vegetation in the study areas was measured using 
a modified James and Shugart (1970) method. Ten 
0.04 ha circles (22 m diameter) were established in 
each study area, one centered on each of five point 
count stations and one at 50 metres in a random direc- 
tion from that point. Within each circle, species, height 
in meters and diameter at breast height (dbh) in cm for 
all trees larger than 7 cm dbh were recorded. Percent 
basal area for each tree species was calculated using 
dbh values. Basal area more accurately reflected tree 
availability to birds than numbers of trees, due to 
heterogeneity in tree size. 

To check for possible visibility biases in the various 
tree species, simultaneous confidence intervals (Chi- 
squared with Bonferroni confidence intervals) were 
constructed by pooling the number of seconds of ob- 
servation for all bird species (Byers et al. 1984). 
Simultaneous confidence intervals were also used to 
assess whether birds preferred or avoided certain tree 
species. Chi-squared test with Yates correction (Zar 
1996) were used post priori to determine if birds had 
a preferred foraging zone within trees. Foraging diver- 
sity was analyzed using the Shannon Index (Shannon 
and Weaver 1949). To assess niche overlap between 
species, the proportion of observations each bird 
species spent in each of the 15 possible tree zones (9 
zones in deciduous trees) was compared between each 
species pair using Morisita’s Index of overlap (see 
Diamond 1983). With this index, potential niche over- 
lap can vary from | (complete overlap) to 0 (no over- 
lap). Foraging height data were analyzed using a 1- 
way Anova (Proc GLM; SAS Institute 1988), and the 
results subjected to a Tukey’s HSD pair-wise com- 
parison test (with P=0.05). 


Results 
Tree Species Preference 

By basal area, White Spruce, Balsam Fir, Trembling 
Aspen and White Birch comprised 46%, 30%, 20%, 
and 4%, respectively, of the available arboreal habitat. 


2004 CUMMING: HABITAT SEGREGATION AMONG SONGBIRDS 47 


BOCH 
(N=128) 
H=2.25 


@ 10- 14.9% [ 5.9.9% LI 0- 4.9% 


FiGuRE 1. Songbird use of zones in coniferous trees. Shadings indicate frequency of observations within a zone (see 
legend). N= number of observations, H= Shannon diversity index. See Table 4 for species abbreviations. 


48 THE CANADIAN FIELD-NATURALIST 


White Spruce was used more than expected by 
Yellow-rumped (Myrtle) Warblers, (Dendroica coro- 
nata), Golden-crowned (Regulus satrapa) and Ruby- 
crowned (R. calendula) kinglets and in proportion to 
its availability by all other species (Table 1). Balsam 
Fir was used more than expected by Boreal Chicka- 
dees (Poecile hudsonicus) and Blue-headed Vireos 
(Vireo solitarius) but less than expected by Tennessee 
Warblers (Vermivora peregrina). White Birch was 
used more than expected by Tennessee and Magnolia 
(Dendroica magnolia) warblers, while Trembling Aspen 
was used less than expected by Boreal Chickadees, 
Golden-crowned and Ruby-crowned kinglets, Blue- 
headed Vireos, Yellow-rumped and Magnolia war- 
blers (Table 1). Bay-breasted Warblers (Dendroica 
castanea) used all tree species in proportion to their 
availability; however, they did exhibit a tendency to 
prefer White Spruce and avoid White Birch that was 
approaching significance. Blackburnian (D. fusca) and 
Black-throated Green (D. virens) warblers were not 
tested due to insufficient sample size (Brennan and 
Morrison 1990). 


Point counts 

All ten species were found in all three study sites, 
except Black-throated Green Warbler, which only oc- 
curred in one of the three study sites. Average bird 
abundance per point count from most to least abundant 
was; Magnolia Warbler (1.2), Yellow-rumped Warbler 
(0.8), Bay-breasted and Tennessee warblers (0.7), 
Blackburnian Warbler and Ruby-crowned Kinglet (0.4), 
Blue-headed Vireo (0.3), Boreal Chickadee and Gol- 
den-crowned Kinglet (0.2) and Black-throated Green 
Warbler (0.1). 


Foraging Zones 

In coniferous trees the most diversely foraging spe- 
cies (in order) were Yellow-rumped Warblers, Ruby- 
crowned Kinglets, Magnolia Warblers, and Boreal 
Chickadees (Figure 1). These species were observed in 
most tree zones and had the highest diversity indices 
(Shannon and Weaver 1949). The least diversely forag- 
ing species were Black-throated Green Warblers, Black- 
burnian Warblers and Blue-headed Vireos (Figure 1). 


Vol. 118 


In the vertical zones, Ruby-crowned Kinglets, and 
Tennessee and Yellow-rumped warblers used the 
bottom of the tree, less than expected, while Boreal 
Chickadees and Golden-crowned Kinglets used the 
top and bottom less than expected (Table 2). Black- 
burnian and Bay-breasted warblers used the bottom 
two zones less than expected, while Blue-headed Vireos 
used the top two zones less than expected (Table 2). 
For the horizontal zones Bay-breasted Warblers and 
Blue-headed Vireos both used small twigs on the 
ends of branches (zone T) less than expected, while 
Tennessee and Blackburnian warblers used small 
twigs (zone T) more than expected (Table 2). Yellow- 
rumped Warblers use large inner branches (zone B) 
less than expected, while Boreal Chickadees, Golden- 
crowned and Ruby-crowned kinglets and Magnolia 
Warblers showed no significant difference in their use 
of inner and outer branches. Black-throated Green 
Warblers were not tested due to insufficient sample 
size (Brennan and Morrison 1990). In deciduous trees, 
only six of the ten species had a large enough sample 
size to test foraging zone use (Figure 2). In deciduous 
trees, Blackburnian Warblers used the top of the trees 
more than expected. Yellow-rumped, Magnolia and 
Blackburnian warblers used zone B less than expected, 
and Tennessee Warblers used zone T more than ex- 
pected (Table 2). 


Niche Overlap 

In conifers, niche overlap varied from 0% between 
Blue-headed Vireos and Blackburnian Warblers to 
95% between Yellow-rumped Warblers and both 
Magnolia Warblers and Boreal Chickadees The four 
most diversely foraging species, Boreal Chickadees, 
Ruby-crowned Kinglets, and Yellow-rumped and Mag- 
nolia warblers, all overlapped each other by >80%. 
In deciduous trees, the most foraging overlap was 
between Yellow-rumped and Blackburnian warblers 
and between Magnolia and Tennessee warblers; both 
species pairs overlapped by 88% (Table 3). The least 
amount of overlap in deciduous trees (40%), was be- 
tween Black-throated Green and Bay-breasted warblers. 


TABLE |. Bird use of different tree species compared to the tree’s availability (y? test with Bonferroni confidence intervals, 


DF=3 for all species). 


Use 
Bird Species 


Boreal Chickadee 
Golden-crowned Kinglet 


> Expected 


Balsam Fir 
White Spruce 


Ruby-crowned Kinglet White Spruce 
Blue-headed Vireo Balsam Fir 
Tennessee Warbler White Birch 
Magnolia Warbler White Birch 
Yellow-rumped Warbler White Spruce 
Bay-breasted Warbler - 
Significance: *< 0.05, **<0.01, ***<0.001 


< Expected ye Pp 
Trembling Aspen 43.3 ss 
Trembling Aspen 10.1 A 
Trembling Aspen 529 : 
Trembling Aspen 6.3 ‘i 
Balsam Fir 50.7 tte 
Trembling Aspen 10.9 
Trembling Aspen DAD Ee 
= 5.0 n.s. 


2004 CUMMING: HABITAT SEGREGATION AMONG SONGBIRDS 49 
YRWA MAWA 
(N=54) (N=64) 
H=1.85 H=1.85 H=1.90 
BLBW 
(N=53) 
H=1.91 H=1.35 H=1.25 
FiGure 2. Songbird use of zones in deciduous trees. Shadings indicate frequency of observations 
within a zone (see legend). N= number of observations, H= Shannon diversity index. 
See Table 4 for species abbreviations. 
Foraging Height average foraging height of Blackburnian Warblers was 


In coniferous trees, the average foraging height of 
Blackburnian Warblers was significantly higher than 
all other species except Black-throated Green and Bay- 
breasted warblers (Table 4). Black-throated Green 
and Bay-breasted warblers both foraged significantly 
higher than Magnolia Warblers and Blue-headed Vir- 
eos. Foraging heights of all other species were not sig- 
nificantly different (Table 4). In deciduous trees, the 


significantly higher than that of Blue-headed Vireos 
and Magnolia Warblers but not significantly different 
from other species. 


Canopy vs. Subcanopy Use 

Blackburnian, Black-throated Green, and Tennessee 
warblers were seen in canopy level trees (>12 metres) 
in two-thirds or more of all observations (Table 5). At 
the other extreme, two-thirds of the observations on 


50 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE 2. Bird use of tree zones where zone use was significantly more or less than expected (x? test with Yates’s correction; 


DF=!1 for all species). 


Species Horizontal Zones 
T,M,B Use OG 
In Coniferous Trees 
Boreal Chickadee n.s 
Golden-crowned Kinglet ns. 
Ruby-crowned Kinglet ns. 
Blue-headed Vireo at < 20.9 
Tennessee Warbler al > 14.9 
Blackburnian Warbler ay > 18.5 
Magnolia Warbler ns. 
Yellow-rumped Warbler B < 14.6 
Bay-breasted Warbler at < 10.5 
In Deciduous Trees 
Tennessee Warbler T > 30.3 
Blackburnian Warbler B < 24.9 
Magnolia Warbler B < 155 
Yellow-rumped Warbler B < 233) 


Vertical Zones 


IP 1 to5 Use Ye RB 
sD) < 20.4 nce 
We) < 14.5 cae 
5 < 16.5 sats 
a Ie) < 1h oe 
Fata 5 < ig? “ook 
nas 4,5 < 7.4 mee 
3 > 13.7 ie 
cates 3) < 10.1 ar 
ae 4,5 < 7.5 ne 
eae ns. 
se I > 6.6 * 
eae ns. 
bt n.s. 


Significance: *< 0.05, **<0.01, ***<0.001; T = outer part of branch with new growth, fresh needles and small twigs; M = 
middle layer of older needles and larger branches; B = inner part of branch, largely devoid of needles with large diameter 


branches; | to 5: vertical zones, 1 = top, 5 = bottom. 


TABLE 3. Average (mean + SD) foraging height (metres) in 
coniferous and deciduous trees. Species marked with * for- 
aged significantly higher than other species (1-way Anova 
with Tukey’s HSD test, P=0.05). 


Bird Species Coniferous Deciduous 
Boreal Chickadee 6.1 + 3.4 ONS: 2 
Golden-crowned Kinglet TOE 229 ODES al 
Ruby-crowned Kinglet 7.3+44.7 9.5+4.7 
Blue-headed Vireo 6.1+2.0 6.3+3.8 
Tennessee Warbler SiOFE SE 5S) aed] 
Blackburnian Warbler esi Bel /a3) ay Thee Spoke 
Magnolia Warbler DR) Baa 6.9 +5.0 
Black-throated Green Warbler 12.7 + 7.8* 11.44+3.8 
Yellow-rumped Warbler 7.0+4.2 NIMs Site) 
Bay-breasted Warbler 10.3 + 4.8* loves Shs 


Coniferous: Blackburnian Warbler foraged significantly high- 
er than all but Black-throated Green Warbler and Bay-breasted 
Warbler. Black-throated Green Warbler and Bay-breasted War- 
bler foraged significantly higher than Blue-headed Vireo and 
Magnolia Warbler. 

Deciduous: Blackburnian Warbler foraged significantly high- 
er than Blue-headed Vireo and Magnolia Warbler. 


Magnolia Warblers and Boreal Chickadees were in 
subcanopy trees (Table 5). Blue-headed Vireos and 
Ruby-crowned Kinglets were approximately equal in 
their use of canopy and subcanopy trees, while Golden- 
crowned Kinglets and Yellow-rumped and Bay-breast- 
ed warblers tended to be observed slightly more often 
in canopy level trees (Table 5). Yellow-rumped Warblers 
were observed in canopy level trees significantly more 
than Magnolia Warblers (Z=5.2, P<0.001) and Boreal 
Chickadees (Z=3.4, P<0.001). 


Discussion 
Tree Species Preference 

The four main tree species in the study area (White 
Spruce, Balsam Fir, Trembling Aspen, and White 
Birch), were not equally abundant, nor equally large 
(Cumming 1995). If abundance alone had been used 
as the criteria for tree availability for foraging then 
Balsam Fir would have appeared to comprise a larger 
proportion of available habitat than it actually did. 

Patches of old, White Spruce-dominated forest are 
relatively rare in Saskatchewan (Kabzems et al. 1986; 
Acton et al. 1998); therefore, it was expected that 
birds which prefer White Spruce would selectively 
occupy this habitat. The data appear to support this, 
as all species in the study used White Spruce either in 
proportion to, or greater than, its abundance. Yellow- 
rumped Warblers and Golden-crowned and Ruby- 
crowned kinglets all used White Spruce more than 
expected. Boreal Chickadees and Blue-headed Vireos 
used White Spruce in proportion to its abundance and 
used Balsam Fir more than expected. All five species 
used Trembling Aspen less than expected (Table 1). 
This is similar to their behaviour elsewhere, as other 
researchers have found these five species to be conifer 
specialists (Keast and Saunders 1991; Ingold and 
Wallace 1994; Ingold and Galati 1997). 

Trembling Aspen was the least favored tree; it was 
used less than expected by half the bird species in the 
study. Previous research has suggested differences in 
insect abundance may exist between aspen and other 
tree species (Ives and Wong 1988). Holmes and Robin- 
son (1981) suggested structural characteristics of aspen 
trees and their leaves may impede foraging by insec- 
tivorous birds. However, Greenburg (1979) felt that 
Yellow-rumped and Bay-breasted warblers were phys- 


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CUMMING: HABITAT SEGREGATION AMONG SONGBIRDS 


2004 


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Sy THE CANADIAN FIELD-NATURALIST 


TABLE 5. Bird use of canopy versus subcanopy trees, from 
most to least canopy using species. 


Number of Observations 


Bird Species Canopy Subcanopy 
Blackburnian Warbler 62 ii 
Black-throated Green Warbler iL) 5 
Tennessee Warbler 80 41 
Yellow-rumped Warbler 94 63 
Bay-breasted Warbler 73 55 
Golden-crowned Kinglet 16 12 
Ruby-crowned Kinglet Dil, 25 
Blue-headed Vireo 24 25 
Boreal Chickadee 29 50 
Magnolia Warbler 60 126 


ically adapted to foraging in conifers and both their 
body size and foraging habits impeded foraging in 
deciduous trees. 

Tennessee and Magnolia warblers both used White 
Birch more than expected. Morse (1989) felt that 
many songbirds preferred various Betula species for 
mechanical reasons; they had smaller leaves than other 
trees (especially Fagus, Acer and Populus) and were 
easier for gleaning songbirds to reach. Other studies 
have found various species of birch have higher den- 
sities of lepidopteran larvae than other trees (Holmes 
and Robinson 1981; Holmes and Shultz 1986). An 
insufficient sample size prevented testing of Black- 
throated Green Warbler’s tree species preference. It is 
notable, however, that 50% of the foraging observa- 
tions on this species were in White Birch, even though 
these trees represented only 4% of the available habitat. 
Other researchers (Holmes and Robinson 1981; Robi- 
chaud and Villard 1999) have also found that Black- 
throated Green Warblers are partial to birch and use 
it greater than expected by chance alone. 

Tennessee Warblers used Balsam Fir less than ex- 
pected, while Blackburnian and Black-throated Green 
warblers were never observed in this tree. Although 
Blackburnian and Black-throated Green warbler use 
of this tree could not be tested, the fact that Balsam 
Fir comprised 30% of the available trees suggests 
that both birds were avoiding it. Blackburnian and 
Black-throated Green warblers were consistently ob- 
served high in the canopy, so Balsam Fir with an 
average height of 11 + 5.6 metres may have been too 
short to attract these species (Cumming 1995). Branch 
structure may also have played a role in some birds’ 
avoidance of fir. Differences in branch shape have been 
attributed to several bird species’ preference of Red 
Spruce (Picea rubens) over White Spruce (Morse 
1989; Parrish 1995). Balsam Fir branches were less 
ridged than those of White Spruce (personal observa- 
tion). Birds which preferentially foraged on thin outer 
twigs appeared to have more difficulty foraging on the 
tips of Balsam Fir branches than they did in White 


Vol. 118 


Spruce (personal observation). This was particularly 
noticeable in Tennessee Warblers because this species 
focused most of its foraging activities on the outer 
twigs and new growth in both coniferous and decid- 
uous trees. In contrast, birds which foraged in many 
tree zones (Yellow-rumped, Magnolia, and Bay-breast- 
ed warblers) appeared to be less affected by branch 
structure and they used Balsam Fir in proportion to 
its availability. 


Foraging 

Boreal Chickadees were the only permanent resi- 
dents in the study and one of the few passerines that 
are resident in the boreal forest (Ficken et al. 1996). It 
is considered an advantage for birds to be flexible in 
their foraging habits when they live in an unpredic- 
table environment, like the boreal forest (Hunt and 
Flaspohler 1998). Boreal Chickadees were diverse 
foragers, using most tree zones and overlapping more 
than 80% with Ruby-crowned Kinglets and Yellow- 
rumped, Magnolia, Bay-breasted and Tennessee warb- 
lers. Other studies found Boreal Chickadees had an 
average foraging height of five metres, and used all 
parts of the tree branch (Sabo and Holmes 1983; 
Ficken et al. 1996). Similarly, I found Boreal Chicka- 
dees foraged at an average height of six metres and 
showed no significant difference in their use of inner 
and outer branches. They did however, use the top 
and bottom of trees less than expected. Boreal Chick- 
adees may have avoided the tops of trees because this 
zone was heavily used by several other species. Boreal 
Chickadees have been shown elsewhere to shift their 
foraging activity lower in the trees due to competition 
from Black-capped Chickadees (Vassallo and Rice 
1981). Boreal Chickadees also tended to forage in 
small subcanopy trees more than any other species 
except Magnolia Warblers. 

Ruby and Golden-Crowned kinglets were similar 
in their behaviour; however, Ruby-Crowned Kinglets 
were usually higher in the trees and were more diverse 
foragers. Rabenold (1978) found that where Ruby- 
crowned Kinglets and Golden-crowned Kinglets oc- 
curred together, Ruby-crowned Kinglets were more 
diverse in their foraging activity and occurred higher 
in the canopy than Golden-crowned Kinglets. Franz- 
reb (1984) felt this was due to social dominance of 
Ruby-crowned over Golden-crowned kinglets. No 
interspecific social interactions were observed in this 
study; however, Ruby-crowned Kinglets were twice 
as abundant as Golden-crowned Kinglets and may 
have influenced the behaviour of the latter through 
numerical dominance. 

Large inner and middle branches were favoured by 
both Blue-headed Vireos and Bay-breasted Warblers; 
this was an area of the trees that was seldom used by 
other species. There was only moderate foraging over- 
lap (47%) between Blue-headed Vireos and Bay- 
breasted Warblers, as they foraged at significantly dif- 
ferent heights. Foraging on large inner and middle 


2004 


branches is similar to the behaviour these two species 
have displayed elsewhere (Williams 1996; James 1998). 

Both Blackburnian and Black-throated Green war- 
blers foraged high in the trees (significantly higher than 
most other species), and had larger foraging overlap 
with each other (85%) than they did with any other 
species. These warblers appeared to be specialized in 
a tree-top niche, and have been identified in many stud- 
ies as feeding, singing, and nesting high in conifer- 
ous trees (Morse 1993, 1994). Indeed, Morse (1994) 
found that Blackburnian Warblers were unlikely to 
be found in forest without at least some conifers over 
18 metres tall. Unlike MacArthur (1958) and Morse 
(1989), I found no evidence that Black-throated Green 
Warblers were socially dominant over any other spe- 
cies in the study area. This may have been because 
Black-throated Green Warblers were the least abundant 
bird in the study and only occurred in one of three 
study sites. 

Similar to their behaviour elsewhere, Tennessee 
Warblers were observed at a variety of heights in all 
tree species, but foraged mainly on the terminal foli- 
age (Rimmer and McFarland 1998). Foraging on the 
tips of the branches and using many different trees 
may help Tennessee Warblers (a Vermivora warbler) 
fit into a community (spruce-fir forest) that already 
has several coexisting species of Dendroica warblers 
(Morse 1989). Tennessee Warblers also used White 
Birch more than any other species except Magnolia 
and Black-throated Green warblers. Greenburg (1979) 
felt that small warblers were better able to forage on 
deciduous foliage than larger warblers for the physical 
reasons. Tennessee, Magnolia and Black-throated Green 
warblers were smaller than most species, except for 
the two kinglets, and Boreal Chickadees, which were 
conifer specialists. Therefore, they may have been 
using a food resource that the larger Yellow-rumped 
and Bay-breasted warblers would have trouble 
exploiting. 

I found Magnolia Warblers used subcanopy trees 
more than any other species and had the lowest aver- 
age foraging height of all species observed. This was 
similar to Hall (1994), who found Magnolia Warblers 
used mainly young conifers and were seldom found 
at any great height. Magnolia Warblers displayed no 
significant difference in their use of inner, middle and 
outer parts of branches. This was likely because they 
most often foraged young trees; there was far less 
difference in branch structure between inner and outer 
parts of branches in young trees than there was in large 
old trees (personal observations). 

Yellow-rumped Warbler showed the greatest diver- 
sity of foraging habits. They used almost every tree 
zone in both coniferous and deciduous trees. Yellow- 
rumped Warblers have been noted elsewhere as being 
generalists and this flexibility is thought to help them 
withstand adverse conditions better than their con- 
geners (Rabenold 1978; Hunt and Flaspohler 1998). 
Unlike their behaviour elsewhere (MacArthur 1958; 


CUMMING: HABITAT SEGREGATION AMONG SONGBIRDS 53 


Morse 1989), Yellow-rumped Warblers did not use 
mainly the bottom and inner branches. Yellow-rumped 
Warblers occurred most often in the middle and outer 
branches at mid to upper heights in both conifers and 
deciduous trees, the same zone frequented by Black- 
throated Green Warblers in other studies (MacArthur 
1958; Morse 1989). Morse (1971) and Howe (1979) 
felt that the presence or absence of Black-throated 
Green Warblers caused shifts in Yellow-rumped War- 
bler foraging activity. I observed no social interactions 
between Yellow-rumped and Black-throated Green 
warblers, and as Yellow-rumped Warblers were eight 
times more abundant, there was likely little interspe- 
cific competition between these two species. Indeed, 
Yellow-rumped Warblers experienced the most overlap, 
and likely the most interspecific competition, from 
Magnolia Warblers, Boreal Chickadees, and Ruby- 
crowned Kinglets (Table 4 and Figure 1). 


Niche Overlap 

Northern regions, such as the boreal forest, are 
thought to have such large pulses of seasonally avail- 
able food that for songbirds food saturation occurs 
(Rabenold 1983; Wiens 1989). This seasonal pulse of 
food, combined with a high number of neo-tropical 
migrants, allows greater niche overlap between song- 
birds than occurs in other habitats (Rabenold 1978, 
1983; Wiens 1989). Rabenold (1978) found that spruce- 
fir forests in particular had a high number of songbirds 
which were “...broadly overlapping congeners”. 
Although the present study and Rabenold’s were not 
directly comparable, many of the species were the same 
between the two studies and results from this study 
support his theory of generalist stacking. Species with 
the most generalized foraging behaviour had the high- 
est amount of niche overlap, while those with more 
specialized foraging behaviour had little niche overlap. 


Canopy vs. Subcanopy 

Availabilty of canopy versus subcanopy trees was 
not measured; therefore, I could not test whether birds 
were significantly selecting one over the other. At the 
two extremes, however, it was apparent that Black- 
burnian Warblers were using very tall trees almost 
exclusively, while Magnolia Warblers were using most- 
ly young regenerating trees. 

I tested Yellow-rumped Warblers’ use of canopy and 
subcanopy trees against Magnolia Warblers’ and Boreal 
Chickadees’ because they had a 95% foraging overlap 
(in conifers) with both species (83% with Magnolia 
Warbler in deciduous trees). A Z-test (in results) found 
Yellow-rumped Warblers used tall (canopy) trees sig- 
nificantly more than the other two species. I felt this 
was a significant difference between these species, as 
they used the available habitat in a similar way and 
they did not forage at significantly different heights. 
Magnolia and Yellow-rumped warblers were also the 
two most abundant species in study. 

Previous research has shown old-growth boreal 
mixedwood forest such as those studied here to harbor 


54 THE CANADIAN FIELD-NATURALIST 


unique communities not seen in pre-rotation age forest 
(Erskine 1977; Kirk et al. 1996: Cumming and Dia- 
mond 2002). This diverse songbird community appears 
to be caused by an interaction between a structurally 
diverse habitat and a mixture of generalized and specil- 
ized foraging behaviours of the bird species involved. 
Although this study did not examine multiple age class- 
es of forest, it would appear that the community dyna- 
mics seen here may be fairly unique due to canopy gap 
dynamics that tend to be associated with old-growth 
forest (Hunter 1990). This type and age-class of forest 
has not received the attention it deserves from either 
environmental impact studies or long-term manage- 
ment plans for the boreal forest. The scientific commu- 
nity needs to pay greater attention to the contribution 
old-growth boreal mixedwood forest makes to bio- 
diversity on both a regional and national scale. 


Acknowledgments 

This study was carried out in partial fulfillment of 
the requirements for a M.Sc. degree at the University 
of Saskatchewan. Parks Canada allowed field work to 
be carried out in Prince Albert National Park. Funding 
was provided by the Canadian Wildlife Service, Prince 
Albert National Park, Nature Saskatchewan, and the 
Saskatchewan Wildlife Habitat Development Fund. 
Office, laboratory and computer space was provided 
by the Canadian Wildlife Service in Saskatoon. A spe- 
cial thank you to my husband S. L. Van Wilgenburg 
for help with tables and figures and to my supervisor 
A. W. Diamond for advice and support during thesis 
research and preparation. This manuscript was im- 
proved by helpful comments from S. L. Van Wilgen- 
burg, K. A. Hobson, and A. J. Erskine. 


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Weyerhaeuser Canada. 1998. Twenty-year forest man- 
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haeuser Canada and Saskatchewan Environment and 
Resource Management. Prince Albert, Saskatchewan. 


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Received 10 March 2001 
Accepted 20 May 2004 


Effects of Mid-winter Snow Depth on Stand Selection by Wolverines, 
Gulo gulo luscus, in the Boreal Forest 


JONATHAN D. WRIGHT! and JEssICA ERNST 


Ernst Environmental Services, Box 753, Rosebud, Alberta TOJ 2T0 Canada 
Present address: Box 648, East Coulee, Alberta TOJ 1BO Canada 


Wright, Jonathan D., and Jessica Ernst. 2004. Effects of mid-winter snow depth on stand selection by Wolverines, Gulo gulo 
luscus, in the boreal forest. Canadian Field-Naturalist 118(1): 56-60. 


Wolverines (Gulo gulo luscus) in a study area in the boreal upland forests of northwestern Alberta and northeastern British 
Columbia (approximately 57°N) were noted to be limited to upland landscapes, despite abundant food in adjacent lowland 
landscapes. Snow-tracking suggested that the species was selecting for the densest climax conifer stands for travel in search of 
food. It was hypothesized that snow depth was a limiting factor for Wolverines in the boreal forest during midwinter, and that 
they selected for this stand-type because of the buffering effect of this type of canopy on ground snow-depths. A series of 
snow-depth measurements were collected. Snow depths collected along Wolverine trails were very significantly lower than 
random snow depths collected under upland canopy (F = 32.84, df = 1, P << 0.010). There was a significant buffering effect 
on snow depth indicated for upland canopy (F = 11.1, df = 1, P < 0.010), while adjacent lowland canopy had no significant 
buffering effect on snow depth (F = 3.45, df = 1, P > 0.05). Wolverines were hypothesized to be limited to upland landscapes 
in the study area because of the buffering effect on snow-depth of the stand types found there, and not for reasons of food 
availability. Climax conifer stands were interpreted as being of high importance to Wolverine survival during winter. Conser- 
vation implications include the detrimental effect on Wolverine populations likely to result from current timber harvesting 
practices in the boreal forest. 


Key Words: Wolverine, Gulo gulo luscus, snow depth, boreal forest, stand, landscape, climax, buffering effect, timber harvest- 


ing, Alberta, British Columbia. 


The Wolverine, Gulo gulo luscus, remains among the 
least understood of the world’s forest mesocarnivores. 
Recent research focusing on the life history of the 
species in cordilleran landscapes is beginning to fill 
knowledge gaps. However, landscape features docu- 
mented as being of high importance to Wolverines in 
mountainous regions — for example, the alpine zone 
(Eric Lofroth, personal communication [see Acknow- 
ledgments section for affiliation]) — are unavailable to 
forest populations. 

In the boreal forest, Wolverines are believed to be 
declining from densities that appear to be lower than 
in any of the other landscapes in which they occur 
(Banci 1994). As they are less observable in the forest 
due to the unbroken forest canopy, Wolverine study in 
the forest ecoprovinces is an even greater challenge 
than in alpine habitats. It is therefore not surprising that 
Banci (1994) has suggested that stand-level and land- 
scape scale “habitat use by Wolverines in forests has 
not been adequately investigated”, and that research 
is needed to study the “habitat needs of Wolverine in 
forests, because there is no sound basis for developing 
habitat management prescriptions at the stand level”. 
In forested areas of the northwest, the Wolverine is 
the furbearer about which wildlife agencies have the 
greatest concern (Bill Johnson, personal communica- 
tion). 

There has been agreement among researchers that 
Wolverine “habitat is probably best defined in terms 
of adequate year-round food supplies in large, sparsely 


inhabited wilderness areas, rather than in terms of 
particular types of topography or plant associations” 
(Kelsall 1981)*. Seasonal shifts by Wolverine in cor- 
dilleran landscapes from the alpine zone in the summer 
to subalpine forest in the winter have been attributed 
to the availability of food (Banci 1994), or are thought 
to be related to avoidance of high temperatures or 
humans (Hornocker and Hash 1981). While snow 
depth has been investigated as a limiting factor for 
Fishers, Martes pennanti (Krohn et al. 1995; Raine 
1983), it has not been implicated as a factor affecting 
habitat selection in the Wolverine. 

This paper presents the hypothesis that Wolverines in 
the boreal forest are limited by mid-winter snow con- 
ditions, which in turn affects stand, and in this case, 
landscape selection. 


Methods 

Snow tracking is being increasingly recognized as a 
reputable scientific tool in wildlife studies, management 
and conservation, with efforts being made to establish 
standardized terminology, institute university courses 
on the subject, and to establish networks of trackers 
throughout the North American continent (Rezendes 
1999; Zielinski and Kucera 1995). Snow tracking may 
be the only practical way of learning details of Wol- 
verine habits and habitat use, as such details are not 
adequately provided by radio-telemetry studies (John 
Krebs, personal communication; Eric Lofroth, per- 
sonal communication). 


56 


2004 


Wolverine and other furbearer activities were mon- 
itored during three winter seasons (1997-2000) utiliz- 
ing snow-tracking methods. Approximately 34 000 km 
were traveled by truck, snowmobile, cross-country skis 
and on foot, in search of furbearer tracks, with an 
emphasis on locating Wolverines. The tracking efforts 
covered approximately 1100 km/? in the region of the 
border country of Alberta and British Columbia known 
as “Chinchaga” (after the Chinchaga River), at approx- 
imately 57° north latitude. The tracking area offered 
conditions especially conducive to this type of study, 
in that the landscape could be conveniently divided into 
distinct upland and lowland components. These com- 
ponents offered markedly different and readily divis- 
ible stand characteristics as a result of the “Great Chin- 
chaga Fire” which blanketed the area during the 1950s 
(Don Williams, personal communication), burning 
all but some upland ridges. Fifty years following the 
fire, two distinct forest types predominate: early-to-mid 
seral second-growth of predominantly aspen or pine 
in the lowlands, with more limited late-seral, climax 
or “over-mature” stands predominated by White and 
Black spruce (Picea spp.), and mostly dead and dying 
Populus spp. in the uplands. The lowland forest may 
be further characterized as forming a mosaic broken 
by broad expanses of relatively open Black spruce fen 
and willow (Salix sp.) muskeg. The upland forest cover 
is by comparison more continuous. 

The area is thoroughly criss-crossed by open linear 
corridors at varying stages of regeneration that were 
cleared to conduct seismology work, and have been 
implicated in possible predator/prey imbalances. Seis- 
mology work and subsequent oil and gas exploration 
and extraction activities have been traditional in the 
area since the 1950s (Brody 1981). The area is expe- 
riencing a surge of such activities at present, and there 
was virtually no segment of the area covered in track- 
ing Wolverines that was not impacted by these activ- 
ities at some point during the three winter seasons. 

Moose (Alces alces) reach some of their highest 
densities in North America in the area (Brody 1981), 
being most heavily distributed in the lowlands, and 
frequent in the uplands. Groups of Woodland Caribou 
(Rangifer tarandus caribou) are frequent at low den- 
sities in the lowlands. These are the only two com- 
mon ungulate species in the region, and they offer a 
generous prey-base for the area’s healthy Grey Wolf 
(Canis lupus) population, whose kills in turn offer a 
ready food source for the infrequent Wolverines. 

When Wolverine tracks were located, the individuals 
were fore-tracked (older trails) or back-tracked (fresh 
trails) in order to gain insights into important and 
little-known details of the creature’s use of the boreal 
forest, including landscape and stand use. Tracks were 
accessed in the morning by snowmobile or truck, and 
followed on foot as long as daylight permitted, with 
the exception of one overnight excursion conducted in 
order to assess advantages and disadvantages to the 


WRIGHT AND ERNST: STAND SELECTION BY WOLVERINES 57 


tracking process of remaining in situ. Pertinent details 
of the Wolverine’s behavior were logged in field note- 
books during the tracking event as well as being 
photographically documented. 

Snow depth measurements (in centimetres) were 
taken over a time frame of approximately | week, char- 
acterized by below-freezing temperatures, between 
periods of snowfall. Snow depths were taken at loca- 
tions determined using random numbers generated by 
a portable computer (calculator). The number indicated 
the number of paces to be taken on the ground, either 
within a grid-square (random snow depths), or along a 
Wolverine or Lynx (Lynx canadensis) trail, at which 
point a snow-depth was taken. 

Bes following snow-depth data were recorded: 

Ten random depths on open upland sites (to quantify 
“non-buffered” upland snow depth) with a grid-cell size 
of 1 km?; 

e 27 random depths in forested upland sites (“buffered” 
upland depths) with a grid-cell size of 1 km; 

e Ten random depths on open lowland sites (to quantify 
non-buffered lowland snow depth) with a grid-cell size 
of 1 km’; 

e 25 random depths on forested lowland sites (buffered 
lowland depths) with a grid-cell size of 1 km?; 

e 46 random depths along documented Wolverine trails 
at random paced intervals; 

e 29 random depths along Lynx trails (for comparison 
purposes) at random paced intervals. 


The following comparisons were made utilizing 
ANOVA tests of significance: 

e Snow depths along Wolverine trails versus random for- 
ested upland snow depths; 

e Random forested upland snow depths versus random 
open upland snow depths (= “buffering effect” of upland 
canopy on snow depth); 

e Random forested lowland snow depths versus random 
open lowland snow depths (= buffering effect of lowland 
canopy on snow depth): 

e Snow depths along Lynx trails versus random forested 
lowland snow depths (for comparison purposes). 


Crusting of snow typically occurs in the area of this 
study during late winter to early spring, and occasion- 
ally in late fall, depending on temperatures. Crustless 
(powder) conditions are the norm in this region for 
the majority of the snow season. Snow conditions were 
noted as being of a crustless nature during the collec- 
tion of this data. 

Wolverine reactions to linear corridors such as 
access roads and seismic lines were documented as 
encountered, with special attention being paid to the 
effect that snow conditions on the corridors (compact- 
ed versus undisturbed) had on Wolverine response. 

Efforts were made to locate Wolf kills in both 
representative landscapes in order to monitor such 
important food sources for utilization by Wolverines. 


Results 
Wolverine tracks were located on ten occasions over 
three winters (for a mean of one Wolverine track en- 


58 THE CANADIAN FIELD-NATURALIST 


countered per 3400 km of searching). Thirteen sepa- 
rate tracking events resulted, totaling 20 linear km of 
off-corridor (forest) tracking, or 26.6 total off-cor- 
ridor km using Magoun’s (1985) estimate of adding 
33% to arrive at non-linear distance traveled. Wolver- 
ines were tracked for an additional 12.3 km on linear 
corridors offering conditions of compacted snow, for 
a total estimated tracking distance of 38.9 km. 

Seven Wolf kills (Moose) were located, two in the 
upland landscape and five in the lowland landscape. 
These kills were monitored for Wolverine use. An 
additional road-killed Moose in the lowland landscape 
was monitored for Wolverine use. Of these, Wolver- 
ine were documented as utilizing one of the upland 
Wolf kills. 

Despite frequent available food in the lowland land- 
scape, Wolverine tracks were located only in the upland 
landscape. Wolverines were revealed to have remained 
in the uplands throughout the tracking events. 

One Wolverine tracked was noted to have traveled 
through a burned-over upland area regenerating to 
Lodgepole Pine (Pinus contorta). The second-growth 
pine formed a low (approximately three-metre), dense 
canopy. Heavy use of the burn by Snowshoe Hares 
(Lepus americanus), provided compacted trails on which 
the Wolverine traveled through this second-growth. 

Wolverines often encountered linear corridors during 
their travels. Wolverine diverged from their line-of- 
travel under the forest canopy to travel on 100% of the 
linear corridors encountered that offered conditions of 
compacted snow (n=17), for distances ranging from 
3 — 3000 m. Wolverines did not travel on any of the 
corridors encountered that had undisturbed snow cover 
(n = 16), choosing instead the most direct route across 
in 100% of instances noted. 

There were two instances where an individual Wol- 
verine paralleled an undisturbed east-west corridor 
from just within the south-facing edge of the canopy. 
The individual crossed the respective corridors in- 
volved to select the south-facing aspect for travels of 
470 m and 30 m, before diverging deeper into forest. 
Exposure to the sun had acted in concert with the 
buffering effect of the canopy to reduce snow depth 
from 32 cm on the corridor, to 4 — 11 cm under the 
canopy’s edge. 

Snow depths along routes selected for travel by 
Wolverines were significantly less than random snow 
depths under the upland canopy (F = 32.84, df = 1, P 
<< 0.010). 


TABLE 1: Comparison of random snow depths 


Wolverine Trails 


Range 13-47 cm 
Mean 31 cm 
Standard Deviation 9.03 


Vol. 118 


Random upland snow depths under the canopy were 
significantly less than snow depths in open upland areas 
(F = 11.1, df = 1, P< 0.010), whereas random lowland 
snow depths under canopy were not significantly dif- 
ferent from snow depths in open lowland sites (F = 
3.45, df = 1, P > 0.05). 

Snow-depths along routes selected for travel by Lynx 
were not significantly different from random snow 
depths under the lowland canopy, where Lynx data 
were collected (F = 2.86, df = 1, P > 0.05). 

When Wolverine tracks were discovered exiting the 
forest onto access routes, the Wolverines involved had 
invariably been traveling through dense to extremely 
dense coniferous cover. While subsequent travel on 
compacted linear corridors would frequently take a 
Wolverine through a variety of stand types, the Wol- 
verines invariably selected similarly dense coniferous 
cover for re-entry into the forest. Within the forest, 
Wolverines were frequently, even continuously, ob- 
served to alter their direction of travel (n = 29) to select 
for routes that offered an even slightly denser canopy 
cover (increased buffering effect on snow). Of the ran- 
dom upland sites, 70% were estimated to represent a 
similar cover type to that selected by Wolverines for 
travel, as compared to 20% of random lowland sites. 


Discussion 

Popular and even scientific literature of past decades 
reflected the level of misconception surrounding the 
Wolverine. The Wolverine was seen as unique among 
wildlife in being best adapted to survive during the 
harshest months, to the point of finding winter a sea- 
son of ease (Rausch and Pearson 1972). The animal’s 
adaptations to a winter landscape were provided as 
evidence of this: the dense, luxurious coat (which may 
give the illusion of good underlying physical condition 
in winter); the enormous, well-haired “snowshoe” 
feet cited as enabling the animal to coast effortlessly 
on top of deep snow; the preference for the harshest 
wilderness areas. In fact, in an ongoing study in the 
foothills of northern British Columbia, Wolverines in 
winter were often found upon examination to be in a 
state of energetic stress approaching thresholds of star- 
vation (Don Reid, personal communication); a large 
percentage of dissected Wolverines have empty gastro- 
intestinal tracts (Banci 1994). Unlike such sympatric 
predators as Martens (Martes americana), and col- 
oured foxes (Vulpes vulpes), Wolverines are too large 
to survive on small prey (Banci 1994), and must there- 
fore wander widely, as indicated in all studies, in search 


Random Lowland Locations Lynx Trails 
23 — 52 cm 17-54 cm 
40 cm 36 cm 


Wes) 8.2 


2004 


of less reliable yet more bountiful sources of food 
such as large ungulate carrion. 

While recent research is presenting a different pic- 
ture of Wolverines and their habits, apparent contra- 
dictions still abound. One of the more notable of the 
apparent contradictions in Wolverine behavior is ad- 
dressed by this paper: Why do Wolverines select for 
wide-open landscapes in some parts of their range and 
the densest of cover in other areas such as the boreal 
forest? During the tracking of Wolverines in the boreal 
forest at Chinchaga, patterns of behavior became ap- 
parent. The species limited itself to the upland land- 
scape despite abundant food in the lowlands, con- 
tradicting literature which suggested that wilderness 
conditions and a ready supply of food governed the 
Wolverine’s presence or absence, rather than specific 
landscape and stand features (Kelsall 1981*; Banci 
1994). The observation that Wolverines in the study 
area were limiting themselves to a specific landscape 
(at least during winter) was supported by conversa- 
tions with the area’s fur trappers and wildlife agents, 
who noted that: 

(1) over many decades of trapper experience in the Chin- 
chaga area, Wolverines were encountered in heavily 
timbered upland terrain (Les Sharp, personal com- 
munication); 

(2) in a winter trapper’s experience over 13 years on a 
lowland landscape trapline adjacent to uplands where 
Wolverines were tracked during this study, not a single 
Wolverine was caught, nor tracks ever detected (Larry 
Smith, personal communication). 

(3) in other lowland traplines in the area with long tra- 
ditions of use none were recalled to have ever yielded 
Wolverine pelts (Bill Johnson, personal communi- 
cation). 


During the tracking efforts, it became quickly appar- 
ent to me that the Wolverines were continuously sel- 
ecting, where feasible, for the path of least snow cover. 
When faced with even a slight thinning of the forest 
canopy (= deeper snow conditions), the animals were 
observed to have paused, selected the densest alter- 
native timber, and altered their route to follow this 
“path of least resistance” in terms of snow depth. The 
Wolverines would only cross such openings if: 

(1) a compacted Showshoe Hare trail crossed the deeper 

snow, which they would then select; 

(2) alternative routes were not available without extensive 
detour, in which case the Wolverine would cross the 
deeper snow over the shortest straight-line path, usually 
at a walking gait. 


Wolverines in the area seemed to favor a “2x” or 
“3x” lope (Zielinski and Kucera 1995) when less 
impeded by snow, suggesting that this was their most 
efficient gait in terms of energy expenditure versus 
distance traveled. They would immediately switch to a 
less efficient walk, however, when faced with unavoid- 
able stretches of deeper snow. 

In comparison, Lynx were observed not to avoid 
deeper snow, readily crossing open areas of the deepest 
snow conditions, as encountered. This is supported 


WRIGHT AND ERNST: STAND SELECTION BY WOLVERINES 59 


not only by the snow depth data, which showed Lynx 
(in direct contrast with Wolverine) to have selected 
paths of insignificantly less snow depth, but also by the 
fact that Lynx were abundantly found in the lowland 
landscape where the buffering effect of the forest 
canopy was demonstrated to be insignificant. 

Wolverines traveling together did so single-file for 

an estimated 95% of the distance traveled, traveling in 
tandem only along ridges where snow cover was neg- 
ligible. It would be interesting to discover if social travel 
in this species represents a survival strategy, and if the 
individuals involved alternate as lead-individual. 

That Wolverines at Chinchaga were not selecting the 

densest canopy conditions in order to avoid detection by 
man or predators is obvious for the following reasons: 

(1) In 100% of cases, they followed open linear corridors 
offering compacted snow conditions when encountered 
(including winter roads up to 18 m wide; freshly 
opened seismic lines; snowmobile trails; all-terrain 
vehicle tire tracks; wind-swept ice on creek-beds bisect- 
ing open muskeg; compacted otter runways on open 
creek beds) for travel of distances up to 3 km; 

(2) Wolverines are more abundant on the open tundra 
(characterized by hard-crusted snow conditions) than 
they are in the forest (Don Reid, personal communi- 
cation; Banci 1994); 

(3) Wolverines seem to prefer the open alpine zones during 
snow-free seasons in cordilleran landscapes (Whitman 
et al. 1986; Banci 1994; Don Reid, personal com- 
munication; Eric Lofroth, personal communication). 


The hypothesis presented by this paper is that Wol- 
verines in the mid-winter boreal forest are limited by 
snow conditions. Conifer-dominated climax stand types 
offer the highest buffering effect on snow conditions 
underfoot. In this study area, such stand-types are 
only found extensively in upland landscapes, which 
accounts for the absence of Wolverines in the lowlands, 
despite abundant food there. Wolverines travel great 
distances in search of unpredictable food (Banci 1994) 
while under conditions of energetic stress, necessi- 
tating that they pay the strictest attention to energy 
expenditure while traveling. To travel in conditions 
of deep powder (unbuffered) snow is to increase risk 
of starvation, and so such conditions are avoided where 
alternatives exist. While the relatively enormous feet of 
the Wolverine, providing low foot loads of 22 g/cm? 
(Knorre 1959) are of great advantage when crusts begin 
to form (Eric Lofroth, personal communication), they 
have little effect in the midwinter powder snows of 
the northwest characterizing this study, as indicated by 
shifts to an inefficient walking gait in deep, midwin- 
ter snow. It is further suggested here that the size of 
the Wolverine’s feet, rather than being an indication 
of their level of mastery of winter conditions, is in- 
stead just the opposite, an adaptation indicative of their 
extreme level of sensitivity to snow depths as a result 
of specialized foraging strategies combined with rela- 
tively large body size leading to conditions of energetic 
stress. While one could argue that Lynx, with their 
equally large feet and specialized foraging strategies, 


60 THE CANADIAN FIELD-NATURALIST 


should therefore show a similar sensitivity to snow 
depths not indicated in this study, it should be remem- 
bered that the Lynx data herein were collected during 
years of high hare numbers. Perhaps during the low 
hare cycle, Lynx must also become more sensitive to 
snow depths. 


Conclusions 

During midwinter, Wolverines in the boreal upland 
forests of northwestern Alberta and northeastern British 
Columbia demonstrate preferences at the stand level 
and landscape scale that are apparently unrelated to 
food availability, consistently selecting for stand types 
that offer the greatest available buffering effect on 
ground snow depth. 

Wolverines require the type of boreal forest habitat 
considered optimum for Martens, the species with 
which they may be considered most highly sympa- 
tric. However, Wolverines appear to be even more 
dependent on climax conifer forests during winter than 
are Martens. Climax conifer cover appears to be of 
high importance to Wolverines in the boreal forest 
because of the buffering effect this stand type has on 
snow depth. 

Forest practices that remove climax growth may 
create conditions that are not conducive to Wolverine 
conservation. As current logging practices in the boreal 
forest are geared towards precisely this type of stand 
removal, timber harvesting may be a key factor in appa- 
rent population declines. Conditions following removal 
of climax cover may not become favorable again for 
Wolverine habitation for many decades (Don Williams, 
personal communication). The situation may be im- 
proved in the case of immediate regeneration to Pinus 
spp., however, with a minimum estimated lapse of 20 
years (several Wolverine generations) before conditions 
become again conducive to Wolverine travel, if not 
other life history components of this species. Efforts 
should be made to preserve linkages of climax conifer 
cover between more extensive areas of intact climax 
forest habitat. 


Acknowledgments 

Funding, accommodations, and ongoing support for 
this project were provided by Pioneer Natural Res- 
ources Canada Inc., to whom the authors are very 
deeply grateful. We would also like to thank the fol- 
lowing (in order they appear in text) for their per- 
sonal communications: Eric Lofroth, British Columbia 
Ministry of Environment, Lands and Parks, Victoria, 
British Columbia; Bill Johnson, Alberta Environment, 
Peace River, Alberta; John Krebs, Columbia Basin 


Vol. 118 


Fish & Wildlife Compensation Program, Nelson, Bri- 
tish Columbia; Don Williams, Alberta Environment, 
Manning, Alberta; Don Reid, British Columbia Minis- 
try of Environment, Lands and Parks, Smithers, British 
Columbia; and the late Les Sharp, as well as Larry 
Smith, fur-trappers in the Chinchaga region of Alberta. 


Documents Cited (marked with * in text) 

Kelsall, J. P. 1981. Status report on the Wolverine, Gulo 
gulo, in Canada in 1981. (Committee on the Status of 
Endangered Wildlife in Canada (COSEWIC), Ottawa, 
Ontario. 47 pages. 


Literature Cited 

Banci, V. 1994. Wolverine. Pages 99-127 in American marten, 
Fisher, Lynx and Wolverine in the western United States. 
Edited by L. F. Ruggiero, K. B. Aubry, S. W. Buskirk, L. 
J. Lyon and W. J. Zielinski. USDA Forest Service General 
Technical Report RM-254. 

Brody, H. 1981. Maps and dreams; Indians and the British 
Columbia frontier. Douglas & McIntyre. Vancouver, British 
Columbia, and Toronto, Ontario. 

Hornocker, M. G., and H. S. Hash. 1981. Ecology of the 
Wolverine in northwestern Montana. Canadian Journal 
of Zoology 59: 1286 — 1301. 

Knorre, E. P. 1959. Ecology of the elk. Jn Transactions of 
the Pechora-Ilych State Game Preserve by G. A. Novikov. 
Translated by The Canadian Wildlife Service 1966. 3895. 
(7): 324 pages. 

Krohn, W. B., K. D. Elowe, and R. B. Boone. 1995. Relations 
among Fishers, snow and martens: development and eval- 
uation of two hypotheses. The Forestry Chronicle 71(1): 
97-105. 

Magoun, A. J. 1985. Population characteristics, ecology, and 
management of Wolverines in northwestern Alaska. Ph. D. 
thesis, University of Alaska, Fairbanks, Alaska. 197 pages. 

Raine, R. M. 1983. Winter habitat use and responses to 
snow-cover of Fisher (Martes pennanti) and marten (Martes 
americana) in southeastern Manitoba. Canadian Journal 
of Zoology 61: 25-34. 

Rausch, R. L., and A. M. Pearson. 1972. Notes on the 
Wolverine in Alaska and the Yukon Territory. Journal of 
Wildlife Management 36: 249-268. 

Rezendes, P. 1999. Tracking & the art of seeing; how to 
read animal tracks and sign. Firefly Books, Willowdale, 
Ontario. 336 pages. 

Whitman, J. S., W. B Ballard, and C. L. Gardner. 1986. 
Home range and habitat use by Wolverines in south- 
central Alaska. Journal of Wildlife Management 50: 460- 
462. 

Zielinski, W. J., and T. E. Kucera. 1995. American Marten, 
Fisher, Lynx, and Wolverine: survey methods for their 
detection. USDA General Technical Report PSW-GTR- 
lsyie 


Received 19 July 2001 
Accepted 26 May 2004 


Wolverine, Gulo gulo luscus, Resting Sites and Caching Behavior in 
the Boreal Forest 


JONATHAN D. WRIGHT and JESSICA ERNST 


Ernst Environmental Services, Box 753, Rosebud, Alberta TOJ 2T0 Canada 
'Present address: Box 648, East Coulee, Alberta TOJ 1B0 Canada 


Wright, Jonathan D., and Jessica Ernst. 2004. Wolverine, Gulo gulo luscus, resting sites and caching behavior in the boreal 
forest. Canadian Field-Naturalist 118(1): 61-64. 


Wolverine (Gulo gulo luscus) caches and resting sites were examined in a study area in the boreal upland forests of northwestern 
Alberta and northeastern British Columbia (approximately 57°N). Cache sites were in climax, or ““overmature” stands of Black 
Spruce (Picea mariana) or mixed-wood of high complexity, dominated by conifers, and in which the Trembling Aspen (Populus 
tremuloides) and Balsam Poplar (Populus balsamifera) component consisted of mostly dead or dying trees characteristic of 
such old growth in the boreal uplands. Sites offered relatively good visibility of the surrounding stand. Sites were never 
located in the dense to extremely dense homogenous spruce stands documented as being favored for travel by Wolverines in the 
study area. The better used cache complexes were accessed by numerous weil-used trails made by the Wolverines themselves. 
Caches consisted of the bones, hide and hair of Moose (Alces alces) believed to have been killed by Grey Wolves (Canis 
lupus). Caches were classified as “simple caches” composed of a single feeding site and/or excavation and “cache complexes” 
involving one or more feeding “stations”, latrines, resting sites, and climbing trees that may have been used as avenues of escape 
from competitors/predators. Resting sites were located atop the snow in relatively open locations that offered good visibility 
of the surroundings. Climax stands were implicated as being of importance to Wolverine caching behavior. Conservation 
implications include the detrimental effect on Wolverine populations likely to result from current timber harvesting practices 


| 
| 
| 


in the boreal forest. 


Key words: Wolverine, Gulo gulo luscus, cache, resting site, climax, boreal forest, stand, Alberta, British Columbia. 


Knowledge gaps exist involving many aspects of 
Wolverine life history in forests, including caching 
behavior and selection of resting sites (Banci 1994). 
This paper presents observations of these aspects of 
Wolverine activities as documented over three winters 
(1997-2000) of snow-tracking efforts in the boreal 
upland forests of the border between northwestern 
Alberta and northeastern British Columbia, at approxi- 
mately 57°N. 


Methods 

Wolverine and other furbearer activities were moni- 
tored during three winter seasons (1997-2000) utilizing 
snow-tracking methods. The tracking efforts covered 
approximately 1100 km? in the region of the border 
country of Alberta and British Columbia known as 
“Chinchaga” (after the Chinchaga River). Tracking is 
being increasingly recognized as a useful scientific tool 
in wildlife studies, management and conservation, with 
efforts being made to establish standardized termi- 
nology, to institute university courses on the subject, 
and to establish networks of trackers throughout the 
North American continent (Rezendes 1999; Zielinski 
and Kucera 1995). Snow tracking may be the only 
practical way of learning details of Wolverine habits 
and habitat use, as such details are not adequately pro- 
vided by radio-telemetry studies (Eric Lofroth, personal 
communication). 

Approximately 34 000 km were traveled by truck, 
snowmobile, cross-country skis and on foot, in search 


of furbearer tracks over the course of the three winters, 
with an emphasis on locating Wolverines. Markedly 
different and readily divisible stand characteristics are 
a feature of the area as a result of the “Great Chinchaga 
Fire” which blanketed the region during the 1950s 
(Don Williams, personal communication), burning all 
but the upland ridges. Fifty years following the fire, 
two distinct forest types predominate: early-to-mid seral 
second growth of predominantly aspen or pine in the 
lowlands, with more limited late-seral, climax or “over- 
mature” stands predominated by White Spruce and 
Black Spruce (Picea spp.), and mostly dead and dying 
Populus spp. in the uplands. The lowland forest may 
be further characterized as forming a mosaic broken by 
broad expanses of relatively open Black Spruce fen and 
open Willow (Salix spp.) muskeg. The upland forest 
cover is by comparison more continuous. 

Moose (Alces alces) reach some of their highest 
densities in North America in the area (Brody 1981), 
being most heavily distributed in the lowlands, and 
frequent in the uplands. Groups of Woodland Caribou 
(Rangifer tarandus caribou) are frequent at low densi- 
ties in the lowlands. These ungulates offer a generous 
prey base for the area’s healthy Grey Wolf (Canis lupus) 
population, whose kills in turn offer a ready food 
source for the infrequent Wolverines. 

When Wolverine tracks were located, the individ- 
uals were fore-tracked (older trails) or back-tracked 
(fresh trails) in order to gain insights into important 
and little-known details of the creatures’ use of the 


61 


62 THE CANADIAN FIELD-NATURALIST 


boreal forest, including landscape and stand use. Tracks 
were accessed in the morning by snowmobile or truck, 
and followed on foot as long as daylight permitted, 
with the exception of one overnight excursion con- 
ducted in order to assess advantages and disadvan- 
tages to the tracking process of remaining in situ. 
Pertinent details of the Wolverines’ behavior were 
logged in field notebooks during the tracking event 
as well as being photographed. 


Results 

Wolverine tracks were located on ten occasions over 
three winters (for a mean of one Wolverine track en- 
countered per 3400 km of searching). Thirteen sep- 
arate tracking events resulted, each lasting approxi- 
mately six hours, and totaling 20 linear km of off- 
corridor (forest) tracking, or 26.6 total off-corridor 
km using Magoun’s (1985) estimate of adding 33% to 
arrive at non-linear distance traveled. Wolverines were 
tracked for an additional 12.3 km on linear corridors 
offering conditions of compacted snow, for a total 
estimated tracking distance of 38.9 km. Five cache sites 
and three resting sites were encountered and docu- 
mented using a GPS unit, camera, and sketches in 
field notebooks. 

Despite frequent available food in the lowland land- 
scape, Wolverine tracks were located only in the up- 
land landscape. Wolverine were revealed to have re- 
mained in the uplands throughout the tracking events. 

Wolverine often encountered linear corridors during 
their travels. Wolverine diverged from their line of 
travel under the forest canopy to travel on 100% of the 
linear corridors encountered that offered conditions of 
compacted snow (n=17), for distances ranging from 3 
to 3000 m. In one instance, two Wolverines traveling 
together followed a compacted (snowmobile) trail on a 
linear corridor for 1270 m to bring them within 150 m 
of a cache site, at which point they exited the trail at 
right angles to follow the shortest linear distance 
directly to the cache. 

All five caches observed consisted of bones and/or 
sections of the hides of Moose (Alces alces), all of 
which were believed to have stemmed from the Wol- 
verine’s scavenging of Grey Wolf (Canis lupus) kills. 
Bones and hide are believed to be a very important 
component of the Wolverine’s winter economy (Banci 
1994; Haynes 1982). Cache-sites appear to be closely 
frequented until the nutritive value is exhausted, judg- 
ing from the well-used system of trails and tracks of 
varying ages in the vicinity of such sites. 

Cache sites encountered ranged from those consist- 
ing of a single bone process or an excavation (n = 2), 
to those better described as “cache complexes” (n = 3). 
Such complexes were believed to stem from the scay- 
enging of a single kill, resulting in a series of smaller 
caches spread over areas which ranged from highly 
localized to widely radiating. Cache sites were in cli- 
max, or “overmature” stands of Black Spruce (Picea 
mariana) or mixed-wood of high complexity, domi- 


Vol. 118 


nated by conifers in which the Trembling Aspen 
(Populus tremuloides) and Balsam Poplar (Populus 
balsamifera) component consisted of mostly dead or 
dying individuals characteristic of such old growth in 
the boreal uplands. Sites offered relatively good visi- 
bility of the surrounding stand. Sites were never locat- 
ed in the dense to extremely dense homogenous spruce 
stands documented as being favored for travel by 
Wolverines in the study area, perhaps because of the 
limited field of vision in such stands. The better-used 
cache complexes were accessed by numerous well-used 
trails made by the Wolverines themselves, and had 
certain features in common, including: 

e areadily accessible spruce tree (Picea spp.) of rela- 
tively large dbh (diameter at breast height) selected 
and well-used for climbing (perhaps as refuge from 
Wolves, or a safe area to digest between feedings); 

° aresting site characterized as a depression in the snow 
formed by the Wolverine’s body and with a good view 
of surroundings; 

e one or more latrines — specific off-trail sites the Wol- 
verine visited to eliminate wastes; 

e aradiating series of “feeding areas” characterized by 
areas approximately 1.5 m* of well-compacted snow, 
on which bones and bone fragments or sections of 
Moose-hide were found, but not both, and distinguish- 
able from a kill site by the lack of Moose stomach- 
content remains; 

e excavations [often in the snow, but on more than one 
occasion (n = 3) in the earth beneath the overhanging 
boughs of a very large spruce tree], that likely had 
contained fragments of cached food. 


A detailed description of the cache and resting sites 
presented on a site-by-site basis follows, in order as 
encountered. 


Cache #1 

Cache # I was a widely radiating cache complex. 
The original Wolf kill site was believed to have oc- 
curred on a linear corridor regenerating to Lodgepole 
Pine (Pinus contorta) located at 57° 27° 20” N and 
120° 08° 26” W. Old Wolf tracks lead to and from 
this central location, and Wolverine trails of varying 
ages, all apparently more recent than the Wolf tracks, 
were present. Shards of Moose bone were located at 
a feeding area here. 

Three additional caches connected by Wolverine 
trails and radiating from the kill site combined to 
comprise the cache complex. 

(1) At 57° 27° 20” N and 120° 08’ 18” W a cache con- 
sisting of Moose hide remains and an abundance of 
Moose hair was located. 

(2) At 57° 27’ 17” N and 120° 08’ 18” W a fresh excava- 
tion in the earth under the boughs of a very large spruce 
tree was located. This cache was believed to have been 
visited by a Wolverine within several hours prior of 
its discovery. 

(3) At 57° 27° 22” N and 120° 08’ 06” W a cache consist- 
ing of bone shards and fragments was located. The 
shards were larger than those examined at the kill site. 
Further components of this cache included an excava- 
tion in the snow down to the sphagnum ground-cover 
and a latrine. 


2004 


This cache complex was not characterized by a 
climbing tree, which was a notable component of some 
other cache sites. The Wolverine tracked to this cache 
complex was dragging what was believed to be a trap 
by one of its left feet. This hindrance apparently ren- 
dered it incapable of climbing, as nowhere during the 
tracking event was it observed to have done so, 
whereas other Wolverines tracked climbed regularly. 
The trap appeared to affect the individual’s locomo- 
tion as well; unlike other Wolverines tracked, this in- 
dividual never varied its gait from a “2x lope” 
(Zielinski and Kucera 1995). 


Cache # IT 

Cache # II was located 150 m from a linear corridor 
with compacted access (snowmobile trail) that was 
followed by two Wolverines traveling together, as 
mentioned, for 1270 m, before exiting the corridor at 
a right-angle to head directly to the cache. 

This cache is best described as a localized complex, 
and appeared to represent the richest of the observed 
caches as evidenced by heavy use. It was located in a 
climax stand of somewhat stunted Black Spruce. Four 
feeding stations were located here, characterized by 
the existence of bone shards and larger fragments and 
processes of bone. There were three latrines, two con- 
taining white scats and one containing brown scats. 


Resting Site # 1 

A resting site, slightly elevated, was located at the 
north edge of the complex. The resting site consisted 
of a 43 x 35 cm oval depression atop the snow, at the 
base of the remains of a short (30 cm), well-wea- 
thered spruce stump. Snow in the depression was well- 
compacted. The resting site was relatively open, away 
from the boles of living trees, and offered a good 
view of the cache complex and its approaches. It was 
accessed by four well-used trails, at approximately the 
four points of the compass. A few guard hairs were 
present in the depression, along with a few dribbles 
of urine at the edge of the east access. 

A climbing tree (Black Spruce) of approximately 
14 m was located in a dense clump of smaller spruce 
at the southeast edge of the cache complex. The tree 
was larger than the stand’s average and heavily used, 
as evidenced by claw-marks on the trunk, heavy debris 
of bark atop the snow at the tree’s base, and the degree 
of trampling of snow within the spruce clump. Climb- 
ing on the tree appeared to terminate at a point near its 
apex where there was a “witches’ broom” formation 
estimated to be large enough to support a resting Wol- 
verine; it may have been an additional resting site. No 
additional trees in the area of the cache complex bore 
any evidence of having been climbed by a Wolverine. 


Resting Site # 2 

A second resting site was discovered by following 
the trail of the two Wolverines after leaving Cache # 2. 
Approximately 200 m from the cache, this resting site 
again consisted of an oval depression (58 x 43 cm) 


WRIGHT AND ERNST: WOLVERINE RESTING SITES AND CACHING 63 


atop the snow at the base of a relatively large Tama- 
rack (Larix laricina) tree. Snow in the depression was 
well-compacted. This resting site was characterized 
by having more canopy cover than the previous site, 
although the cover was by no means dense. There 
was no cache in the immediate vicinity of this resting 
site, and no apparent climbing tree. 


Cache # III 

Cache # III was located at 57° 26’ 84” N and 
120° 07° 69” W and is best termed a “simple cache”, 
rather than a cache-complex. This cache consisted of a 
single large bone-process and a well-gnawed Moose’s 
hoof. The cache was located by following the previ- 
ous two Wolverines, which evidently were aware of, 
or scented, this cache, as evidenced by their diverting 
their course of travel by approximately 50 m to reach 
the cache. There were no other features characteristic 
of other caches present at Cache # III. 


Cache # IV 

Cache # IV located at 57° 27’ 25” N and 120° 06’ 
81” W is best termed a cache-complex, although not 
extensive. Certain features lacking at the last (simple) 
cache were present at this one. This cache consisted 
of a collection of bone fragments located on a single 
feeding platform under a relatively open canopy in a 
high-complexity climax mixed-wood stand. The cache 
was accessed by numerous well-used Wolverine trails. 
No latrines were discovered at this cache. 

Approximately 10 m from the main cache was an 
excavation in the earth under the overhanging boughs 
of a very large spruce tree, accessed by a Wolverine 
trail, and virtually identical to the excavation discoy- 
ered at Cache # I. This excavation, too, was deemed 
to have contained cached food that had been recently 
recovered by a Wolverine. 

Cache # IV was further characterized by a well-used 
climbing tree (spruce) of larger than average dbh, 
located approximately 4 m from the feeding-station. 
This cache was located approximately 10 m from a 
linear corridor with compacted access (snowmobile 
trail) that was used as a conduit for 700 m in a nor- 
therly direction by the Wolverine pair upon exiting 
the cache. This cache had been visited recently by a 
Marten (Martes americana). 


Cache #V 

Cache # V was located in very large, climax mixed- 
wood dominated by spruce, near the base of a ridge 
adjacent to Resting Site # III. This was a simple cache, 
consisting of a single fresh excavation in the earth 
beneath the overhanging boughs of a large spruce, 
much as described for caches I and IV. The excava- 
tion was also believed to be for the purpose of recov- 
ering cached food. 


Resting Site # 3 

Resting Site # 3 was located along the Wolverine 
pair’s trail at 57° 27° 72” N and 120° 05’ 11” W. As 
in the case of the other resting sites observed, this 


64 THE CANADIAN FIELD-NATURALIST 


one consisted of an oval depression atop the snow 
approximately the same dimensions as the last pre- 
vious two sites (measurements not taken due to this 
similarity). The site was mid-slope on a prominent 
ridge covered by a stand of very large mixed-wood 
dominated by spruce. The site was relatively open, 
and offered good view down and along-slope, and only 
slightly less so upslope. Within 1 m of the resting site 
was a medium-sized pine tree which had been well- 
climbed to a height of approximately 4 m. 

Tracks indicated that this resting site may have 
been utilized by a third Wolverine which was travel- 
ing down slope. It appeared as though this third 
Wolverine left the resting site to travel approximately 
15 m along-slope to meet the Wolverine pair as they 
traveled upslope. The single Wolverine may have util- 
ized the climbing-tree to observe the approaching 
pair. A confusion of tracks resulted where the three 
Wolverines apparently met. It is believed that all 
three animals subsequently traveled upslope together. 


Discussion 

Caches were located in climax stands of less density 
than those apparently favored for travel by Wolverines 
in the study area. Visibility of approaching competi- 
tors/predators (Wolves) may be the deciding factor in 
both cache-site and above-snow resting-site selection. 
That the caches were all located in climax timber re- 
inforces the importance of this stand-type to Wolver- 
ine. Present forest practices that result in the harvest- 


Vol. 118 


ing of climax stands may negatively impact Wolver- 
ine populations. 


Acknowledgments 

Funding, accommodations and ongoing support for 
this project were provided by Pioneer Natural Res- 
ources Canada Inc., to whom the authors are deeply 
grateful. 


Literature Cited 

Banci, V. 1994. Wolverine. Pages 99-127 in American marten, 
Fisher, Lynx and Wolverine in the western United States. 
Edited by L. F. Ruggiero, K. B. Aubry, S. W. Buskirk, L. 
J. Lyon and W. J. Zielinski. USDA Forest Service Gen- 
eral Technical Report RM-254. 

Brody, H. 1981. Maps and dreams; Indians and the British 
Columbia frontier. Douglas & McIntyre. Vancouver, Bri- 
tish Columbia; Toronto, Ontario. 

Haynes, G. 1982. Utilization and skeletal disturbances of 
North American prey carcasses. Arctic 35: 266-281. 

Magoun, A. J. 1985. Population characteristics, ecology, and 
management of Wolverines in northwestern Alaska. Ph.D. 
thesis. University of Alaska, Fairbanks, Alaska, 197 pages. 

Rezendes, P. 1999. Tracking & the art of seeing; how to read 
animal tracks and sign. Firefly Books, Willowdale, Ontario. 
336 pages. 

Zielinski, W. J. and T. E. Kucera. 1995. American marten, 
Fisher, Lynx, and Wolverine: survey methods for their 
detection. USDA General Technical Report PSW-GTR- 
ilyt/s 


Received 19 July 2001 
Accepted 26 May 2004 


Seasonal Home Ranges of Raccoons, Procyon lotor, Using a Common 
Feeding Site in Rural Eastern Ontario: Rabies Management Implications 


SARAH C. ToTTon,! RICHARD C. ROSATTE,? ROWLAND R. TINLINE,? and LAURA L. BIGLER‘ 


'340 Second Avenue West, Owen Sound, Ontario N4K 4L7 Canada 

? Ontario Ministry of Natural Resources, P.O. Box 4840, Peterborough, Ontario K9J 8N8 Canada 

> Geographic Information Systems Laboratory, Queen’s University, Kingston, Ontario K7L 3N6 Canada 

+ Zoonotic Disease Section, Diagnostic Laboratory, College of Veterinary Medicine at Cornell University, P.O. Box 5786, 
Upper Tower Road, Ithaca, New York 14852-5786 USA 


Totton, Sarah C., Richard C. Rosatte, Roland R. Tinline, and Laura L. Bigler. 2004. Seasonal home ranges of Raccoons, 
Procyon lotor, using a common feeding site in rural eastern Ontario: Rabies management implications. Canadian 
Field-Naturalist 118(1): 65-71. 


Thirteen adult Raccoons (Procyon lotor) (six females, seven males) that fed at a garbage dump north of Kingston, Ontario 
were radio-tracked from 21 June to 16 October 1995 to assess their seasonal home ranges and movements. Average 
Minimum Convex Polygon (MCP) summer and fall home ranges for the collared Raccoons were 78.4 ha (SD=46.2 ha) and 
45.6 ha (SD=29.7 ha), respectively. Average grid cell summer and fall home ranges for the collared Raccoons were 143.3 ha 
(SD=40.0 ha) and 116.9 ha (SD=24.9 ha), respectively. Summer ranges of the Raccoons were significantly larger than fall 
ranges using both the MCP method (P=0.05) and the grid cell method (P=0.073). Yearling Raccoons travelled an average 
summer maximum distance from the dump of 2608 m (SD=1964, n=3), more than double the distance of adults (22 yr) at 
1239 m (SD=547, n=10). The population density for the study area in late August 1995 was estimated at | Raccoon/12 ha 
based on an effective area surrounding the dump of 234 ha. Home range and movement data may be useful to design a 
strategy to control Raccoon rabies in Ontario. 


Key Words: Raccoon, Procyon lotor, rabies, communal feeding, disease transmission, field study, home range, telemetry, Ontario. 


Raccoon (Procyon lotor) rabies was first reported 
in Ontario, Canada, during July 1999 (Wandeler and 
Salsberg 1999; Rosatte et al. 2001). Point infection 
control methodologies are currently being used (since 
1999) in Ontario to contain the outbreak to a small 
area (Rosatte et al. 2001). As well, a Raccoon rabies 
model is being developed in Ontario to assist with the 
control of the disease by predicting the rate of spread of 
Raccoon rabies both temporally and spatially. Know- 
ledge on Raccoon home ranges, movements and popu- 
lation dynamics in Ontario is needed to develop and 
validate the rabies model so that it reflects the actual 
sequence of events that occur during a Raccoon 
rabies epizootic/enzootic (Broadfoot et al. 2001). 

The spatial distribution of food is known to influ- 
ence contact rates in Raccoons (Procyon lotor) (Sei- 
densticker et al., 1988). Clumped food resources such 
as garbage dumps may increase potential contact rates 
of Raccoons as these resources cause members of a 
population to congregate from a wide area. This in 
turn may influence rabies transmission in the popula- 
tion (Seidensticker et al. 1988). Range of movements 
of Raccoons using common feeding sites may be a 
useful indicator of potential rabies spread. Such infor- 
mation can be used to design effective baiting strate- 
gies to vaccinate these animals against rabies as well 
as provide input for the development of rabies models. 

Home ranges of Raccoons tend to shift due to sea- 
sonal changes in behavior and therefore must be calcu- 
lated separately for each season (Kauffmann 1982). 


Summer is the family rearing period when lactating 
mothers and their offspring travel together and it is also 
the main dispersal period for yearling males (Mech et 
al. 1968; Fritzell 1978). Fall is a time when the juve- 
niles may disperse and Raccoons prepare for the com- 
ing winter dormancy period (Shirer and Fitch 1970). 
In this study, home range was defined using criteria 
of White and Garrott (1990) as the area within which 
the animal normally moved in a specified time frame, 
in this case the summer and fall of 1995. 

In this study, movements of Raccoons using a com- 
mon feeding site in rural eastern Ontario, Canada, were 
determined by radio-telemetry to assess the size of their 
summer and fall home ranges. The same Raccoons on 
which contact data were obtained in the Totton et al. 
(2002) study were used. In addition, two Raccoons, 
caught at a smaller feeding site (compost heap) were 
tracked periodically to determine their daytime rest- 
ing sites. The population density of Raccoons in this 
study was also measured as it influences home range 
and contact rate. 


Study Area and Methods 

Trapping took place at a private garbage dump 
(44°34’N, 76°20’W) and at a compost bin on the 
grounds of the Queen’s University Biological Station 
40 km north of Kingston, Ontario (44° 35’N, 76° 19° W). 
The area surrounding the dump consisted of farm land 
(livestock), forest, marsh, and cottages (most of which 
were only occupied during the summer). The entire 


65 


66 THE CANADIAN FIELD-NATURALIST 


study area was about 460 ha. Fifteen Raccoons (eight 
females and seven males) were collared between 20 
May and 27 June 1995. Raccoons were captured us- 
ing Tomahawk #106 (Tomahawk Live-trap Company, 
Tomahawk, Wisconsin, USA), and Havahart #1079 
(Havahart Live Trap Company, Niagara Falls, Ontario, 
Canada) live-traps. All Raccoons were ear-tagged 
(numbered size | and 2, National Band and Tag Com- 
pany, Newport, Kentucky), vaccinated against rabies 
(Imrab® inactivated rabies vaccine, Merieux, Inc., 
Athens, Georgia, USA) and canine distemper (Fromm 
D, modified live virus, SOLVAY animal health, Inc., 
Mendota Heights, Minnesota, USA). They were im- 
mobilized by intramuscular injection of ketamine hy- 
drochloride (Rogar/STB Inc., London, Ontario, Can- 
ada) and xylazine hydrochloride [Rompun] (Bayvet, 
Rexdale, Ontario, Canada) [30 mg/kg body weight 
ketamine, 10:1 ratio ketamine:rompun]. We deter- 
mined their sex and extracted a first premolar tooth for 
age determination by cementum analysis (Johnston 
et al. 1987). Each animal was then fitted with an adjus- 
table radio-collar [151 to 152-Mhz] (Lotek Engi- 
neering Inc., Newmarket, Ontario, Canada) and rel- 
eased at its point of capture. 

The radio-tracking system consisted of a four- 
element Yagi antenna, 151 MHz (FM) transmitters 
mounted on whip antenna collars (Lotek Engineering, 
Newmarket, Ontario), one programmable hand-held 
receiver (Lotek model SRX-400; Lotek Engineering, 
Inc., Newmarket, Ontario) that operated in the 151- 
152 MHz range, one hand-held compass, and a four- 
wheel drive pick-up truck. Animals were given at 
least seven days to acclimatize to their collars before 
radio-tracking began, in accordance with White and 
Garrott’s (1990) recommendations. The tracking per- 
iod lasted from 21 June to 16 October 1995 with 
attempts being made to locate each Raccoon two to 
three times per week. Since only one telemetry receiver 
was available, sequential rather than simultaneous 
bearings had to be taken. A maximum interval between 
first and last bearings of 10 min was set to minimize 
telemetry error caused by animal movement (except 
for bearings taken during the day when the animals 
were inactive, at which time the interval may have 
been longer). In a study by Gert and Fritzell (1996), 
23% of the locations came from triangulations with 
between-bearing intervals in excess of 8 min. For this 
reason, the 10 min cut-off was deemed reasonable for 
this study. 

Continuous radio-tracking (i.e., location of the ani- 
mals at least every 15 min (Harris et al. 1990)) was 
not feasible with only one receiver; therefore, for this 
study, discontinuous tracking was performed. Location 
estimates were made for each animal three or four times 
between dusk and dawn at roughly 2-h intervals, and 
once during the following afternoon to determine day- 
time resting sites. The tracking schedule was construct- 
ed by randomly selecting six of the collared animals 


Vol. 118 


trapped at the dump for one given tracking night. The 
remaining Raccoons were then tracked on the next 
scheduled night. A different set of Raccoons was ran- 
domly chosen for the following tracking night and so 
on. Dates of tracking nights were randomly selected 
for each week. 

Three types of location estimate techniques were 
used: scanning, triangulation, and homing. Scanning 
involved tuning into the collar frequencies of the dump 
animals while the researcher sat in the middle of the 
dump area. Data from telemetry accuracy tests indi- 
cated a mean transmitter-receiver distance of 240 m+ 
30 m (n=20) when the signal was picked up at a gain 
of 10. A Raccoon was therefore considered to be in a 
radius of this distance from the center of the dump area 
if its signal was detected from there at a gain of 10 or 
less. Most of the locations for the dump animals were 
obtained by triangulation. This technique involves tak- 
ing directional bearings from two to three different 
receiver sites at known locations and using these to 
estimate the true location of a remote transmitter on 
the animal’s collar (White and Garrott 1990). 

Accuracy tests were performed to determine the 
error associated with locations estimated by triangu- 
lation in this study. Bearing accuracy has two compo- 
nents: bias (the average difference between the true 
bearing and the bearing estimated by the receiving 
system for a series of receiver-transmitter locations), 
and precision, which is the standard deviation of these 
errors (White and Garrott 1990). The bias was 9° and 
was significantly different from O° (t=3.61, n=63, 
P<0.001; one sample t-test (Zar 1996)). Therefore 9° 
was subtracted from all bearings. Precision of the sys- 
tem was + 20°. 

Screening criteria derived from telemetry accuracy 
tests were applied to all bearings to eliminate errors 
due to signal bounce. All bearings which did not inter- 
sect with other bearings taken on the same transmit- 
ter were removed. All bearings taken from receiver 
sites associated with large errors (high levels of sig- 
nal bounce) were removed from subsequent analysis. 
Also, bearings taken when the transmitter-receiver dis- 
tance was >2000 m were removed. In addition, all loca- 
tions involving distances between transmitter and rec- 
eiver of over | km were inspected for plausibility. 

A computer program called TRIANG was used to 
estimate animal locations from triangulated bearing 
pairs and to calculate the distance between receiver 
and transmitter for each bearing. TRIANG did not 
compute animal locations when three bearings were 
taken. In this case, locations were determined by plot- 
ting the bearing angles in AutoCAD and estimating 
the centre of the triangle created by the intersection 
of the bearings. 

Locations for two of the collared Raccoons were 
usually obtained by homing (White and Garrott 1990). 
Locations of animals determined by homing are not 
affected by error in the telemetry system; however, they 


2004 


are affected by the researcher’s ability to pinpoint the 
den location on a map (White and Garrott 1990). All 
locations determined by homing in our study were 
plotted by hand on 1:10 000 maps of the area to +50 m 
to obtain Universal Trans Mercator Co-ordinates 
(UTMC). 

Home ranges were estimated using the minimum 
convex polygon (MCP) method (Mohr 1947), since 
this is the only home range method that is strictly com- 
parable between studies (Harris et al. 1990). Ranges 
for both seasons were combined to compare degree of 
overlap between seasons. Because it is advantageous 
to use more than one home range estimate technique 
(Voigt and Tinline 1980), the grid cell method of 
home range analysis (Siniff and Tester 1965) was 
also used. Size of the grid cells was chosen to reflect 
radio fix accuracy based on the results of accuracy tests. 
In order to enclose the uncertainty area associated 
with scanning the dump for transmitter signals (and 
this was larger than the area associated with triangu- 
lation) a grid square would have to measure 480 m on 
each side. Therefore this was the size of grid square 
(23 ha) used in estimating grid cell home ranges. The 
grid was oriented by centering a grid square over the 
dump site. 

Summer home ranges were calculated from teleme- 
try data collected in June, July, and August; Septem- 
ber and October fixes were used to calculate fall home 
ranges. Two female Raccoons were excluded from the 
analysis because insufficient locational fixes (<16) 
were acquired during the study. A Mann-Whitney U 
test (Zar 1996) and Statistica Version 6.0 software 
(StatSoft Inc., Tulsa, Oklahoma) were used to test for 
statistical differences in Raccoon home range size. 
Raccoons were grouped into regular and occasional 
dump visitors based on the number of nights each 
animal was seen at the dump out of a total of 35 
observation nights in a concurrent behavioral study 
(Totton et al. 2002). Raccoons were defined as regu- 
lars if they were observed at the dump on >51% of 
all observation nights and occasionals if they were 
seen at the dump on <51% of the observation nights. 
Since only one telemetry receiver was available, and 
hence discontinuous locational fixes had to be ob- 
tained, detailed analysis of the movement patterns of 
raccoons in this study was not possible. 

Trapping to estimate the Raccoon population size 
took place at the dump from 28 August to 19 Sep- 
tember 1995. At this time of year, juveniles are larger 
and easier to trap and handle than they are earlier in 
the summer (Seidensticker et al. 1988). The number 
of Raccoons in the study area was estimated using a 
modified Petersen Index (Begon 1979). Density of 
the dump population was not based solely on the area 
of the trapping site (garbage dump = 2.3 ha) because 
it was evident from telemetry data that some Rac- 
coons were travelling from a much wider area to feed 
at the dump. Instead, using the number of Raccoons 


TOTTON, ROSATTE, TINLINE, 


AND BIGLER: RACCOONS 67 
calculated by the methods above, estimates were made 
of the crude density of the population as defined by 


Seidensticker et al. (1988) based on the average maxi- 
mum width of the dump Raccoons’ summer (MCP) 
home ranges (1530 m). Since Seidensticker et al. 
(1988) did not specify how this distance was used to 
calculate overall area, a square was centred over the 
dump with each side equal to the distance calculated 
and this was used for calculation of crude density. Its 
area was 234 ha. 


Results 

For locations determined by triangulation, distance 
between the observer and estimated transmitter loca- 
tion ranged from 8.3 m to 1837.9 m and averaged 
330.5 m (SD=239.3 m). Uncertainties in Raccoon 
position for triangulated bearings (tan 20° x trans- 
receiver distance) ranged from +3.0 m to +668.9 m 
with a mean of +120.3 m and standard deviation (SD) 
of 87.1 m (n=1181). Overall locational uncertainty, 
including that associated with scanning and homing 
techniques as well as triangulation, was +146.6 m 
(n=1110). 

No statistical differences were detected between 
male and female ranges for values calculated using 
either the MCP method (P=0.78) or the grid cell meth- 
od (P=0.26). Therefore, male and female data were 
pooled to compare summer and fall ranges. As well, 
only three yearlings were trapped at the dump in this 
study and home range data were available for two of 
those. Consequently, statistical comparisons between 
adult and yearling home range sizes were not per- 
formed. No differences could be detected between 
home ranges of Raccoons that regularly versus occa- 
sionally visited the dump. Average Minimum Convex 
Polygon (MCP) summer and fall home ranges for the 
collared raccoons were 78.4 ha (SD=46.2 ha) and 
45.6 ha (SD=29.7 ha), respectively (Table 1). Average 
MCP summer/fall home range overlap was 31.2 ha 
(SD=17.2). Average grid cell summer and fall home 
ranges for the collared Raccoons were 143.3 ha 
(SD=40.0 ha) and 116.9 ha (SD=24.9 ha), respective- 
ly (Table 1). Average grid cell home range overlap 
between summer and fall was 95.8 ha (SD=27.5). 
Summer ranges of the Raccoons were significantly 
larger than fall ranges using both the MCP method 
(P=0.05) and the grid cell method (P=0.073). 

The most widely ranging Raccoon (a male year- 
ling) in the study was originally trapped at the dump 
on 27 June. It was later located by telemetry near a 
farmhouse 4 km northeast of the dump on 16 July. 
By 18 July, it was visually identified at the dump site 
again where it remained until 8 August. The only other 
yearling of the dump Raccoons wandered a maximum 
of 769 m from the dump during the summer. The aver- 
age distance of the farthest fix from the dump during 
the summer for the yearling Raccoons was 2608 m 
(SD=1964, n=3) and for the adults (2 2 yr), 1239 m 


68 THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE |. Summer (June to August) and fall (September to October) home ranges of 13 Raccoons which fed at a rural 


Ontario garbage dump and at a compost heap in 1995!. 


Method Number Summer 
of fixes range 
(summer) (ha) 
mean (SD)? mean (SD) 

MCP SZe (ile) 78.4 (46.2) 

Grid Cell 44 (6.7) 143.3 (40.0) 


Number Fall Area of overlap 
of fixes range of ranges 
(fall) (ha) 
mean (SD) mean (SD) mean (SD) 
34.1 (9.6) 45.6 (29.7) Sil= (GU) 
33a (92) 116.9 (24.9) 95.8 (27-5) 


'n=13 raccoons {7 males — 6 adults and | yearling; 6 females — 5 adults (all lactating) and 1 yearling} 


2 §D=Standard Deviation 


TABLE 2. Home range, movements and density of Raccoons in different areas of North America 


Location Habitat Home Range Movements 
(km?) (km) 
Ontario rural 0.5-4.0 4-45 
North Dakota rural 0.2-49 1-24 
Minnesota rural 7-12 >3 
Wisconsin 
Toronto urban 0.4 <l 
Ohio urban <0.2 0.4-0.5 


(SD=547, n=10) (P=0.31). The Petersen estimate of 
the number of Raccoons using the dump was 19, 
with upper and lower 95% confidence limits of 35 


and 12 respectively. Crude density was estimated at | 
Raccoon/12 ha. 


Discussion 

Home ranges of Raccoons in North America are 
variable (Table 2) but tend to average around 100 to 
300 ha (1-3 km?) (Kauffmann 1982), but may range 
from 18 to 2560 ha (0.18-25.6 km?) for adult males 
in North Dakota, or 5.1 to 372 ha (0.05-3.72 km?) for 
adult females (Stuewer 1943; Hoffmann and Gotts- 
chang 1977; Fritzell 1978). Urban Raccoons tend to 
have smaller home ranges than rural Raccoons (Ros- 
atte et al. 1991) (Table 2). Range size may also vary 
with season, data collection methods and method of 
home range estimation (Harris et al. 1990). Distribu- 
tion and abundance of food also affect home range 
size (Hoffmann and Gottschang 1977). Small home 
ranges are associated with high population densities 
and abundant food (Hoffmann and Gottschang 1977). 
Fall home ranges of Raccoons in this study were found 
to be significantly smaller than ranges during the sum- 
mer. This was expected as, during the fall, in northern 
areas such as Ontario, Raccoon movement is thought 
to decrease to conserve energy in preparation for the 
winter denning period (Rosatte 2000). Large move- 
ments by Raccoons during the fall, when food sources 
are not as abundant, would likely result in a net ener- 
gy loss thereby decreasing the condition of the animal 


Density Reference 
(/km?) 
4-11 Rosatte and MacInnes 1989 
Rosatte 2000; Rosatte et al. 2001 
0.5-1 Fritzell 1978; Greenwood 1982 


2-6 Schnell 1970; Mech et al. 1968 
Dorney 1954; Schneider et al. 1971 
Rosatte et al. 1991; Rosatte 2000 
Cauley 1970; Schinner 1969 
Hoffmann and Gottschang 1977 


7-85 
45-100 


and decreasing the probability of surviving a harsh 
winter. Movements of Raccoons in this study were 
smaller than noted in other rural Ontario studies and 
in other North American jurisdictions (Rosatte 2000; 
Table 2). That may have been due to the concentrated 
food sources in the vicinity of the dump in this study. 
Raccoons have been known to change their movements 
and home ranges to include new concentrated sources 
of food; Seidensticker et al. (1988) discovered that 
within 27 days, 21 of 23 collared Raccoons living in 
the area had visited their artificial feeding station at 
least once. The feeding site in their rural study area 
was 0.5 ha in diameter and attracted Raccoons from a 
127 ha area. 

Home ranges of male Raccoons tend to be larger 
than those of females (Stuewer 1943: Fritzell 1978). 
The explanation as to why no difference was found 
in this study may have been because either there was 
no difference (due to abundant food sources at the 
dump), or the small sample size prevented the differ- 
ence from being detected. Lack of a difference between 
home ranges of regular and occasional dump visitors 
in this study may have been the result of either a true 
lack of difference or the small sample size involved. 
It may also be that the definition for “occasional” 
was not adequate for comparative purposes. 

The coarseness of the grid used for grid cell home 
range analysis in this study may have led to an over- 
estimate of home range sizes based on the fixes ob- 
tained in this study (White and Garrott 1990) and 
would explain why the grid home range estimates were 


2004 


larger than the MCP estimates. The advantage of the 
grid cell method over the MCP method is that the 
grid method takes into account the precision of the 
telemetry system (White and Garrott 1990). Rac- 
coons in Hoffmann and Gottschang’s (1977) study in 
suburban Ohio with a Raccoon density of 1/1.46 ha 
had average home ranges of 5.1 ha. Fritzell’s (1978) 
study with a very low density of Raccoons in the spring 
and summer averaged 2560 ha. Home ranges of Rac- 
coons in previous Ontario studies (densities of 4 — 
94/km7), varied between 50 and 400 ha (Rosatte 2000; 
Broadfoot et al. 2001). Raccoons in our study fell 
between these two extremes, though their home ranges 
appeared to be smaller than average. However, home 
range estimates obtained from our study data are prob- 
ably underestimates of actual home ranges because the 
study animals were monitored discontinuously over 
only two seasons. In a Niagara-St. Lawrence trap-re- 
capture study, annual movements of Raccoons averaged 
10 km and ranged up to 150 km. In a similar Barrie, 
Ontario, study, nightly movements of 4 km were com- 
mon (Rosatte 1996*; 2000). Unless Raccoons are 
tracked continuously with an accurate system for a 
long period of time (e.g., 1 yr), their movements and 
home ranges will probably be underestimated. 

Radio-telemetry indicated that one yearling male 
Raccoon occupied two distinct and widely separated 
areas during the summer of 1995. The fact that no 
points were located between these two ranges may 
have been because he traversed the distance between 
the two areas in the time between tracking nights. The 
distance between the two discrete areas was 4 km. 
Raccoons have been known to travel this distance in 
a single night (Rosatte 1996*). In addition, the main 
dispersal period for yearling males in some areas is 
May to June (Fritzell 1978) and this might have in- 
clined this Raccoon to make such a movement if he 
were dispersing. A second explanation for the yearling 
male Raccoons’ summer home range pattern is that it 
may have been aided by a vehicle. Raccoons have been 
known to ride on such vehicles as boats, transport 
trucks and rail cars (Rosatte et al. 2001). 

One yearling female Raccoon did not travel as far 
from the dump as the yearling male noted above, in- 
dicating that it may have been a post-disperser, having 
immigrated to the dump area the previous fall. Two 
out of three of the yearlings in this study were found 
in excess of 2 km from their initial capture site within 
the same season of their capture; this indicates that the 
potential for rabies spread in an unvaccinated popu- 
lation using a common feeding site is probably exac- 
erbated by dispersal of the yearlings from the site dur- 
ing the summer. This has been confirmed by Rosatte 
(unpublished data) in a Raccoon rabies epizootic area 
in eastern Ontario. As with the home ranges, men- 
tioned above, these movements are probably minimum 
estimates of actual distances travelled by the dump 
Raccoons. 


TOTTON, ROSATTE, TINLINE, AND BIGLER: RACCOONS 69 


Although caution should be exercised in using data 
from healthy animals to predict the behavior of rabid 
ones, telemetry studies on two other species indicat- 
ed that movements of rabid animals are not very 
different from the movements of healthy ones (Storm 
and Verts 1966; Artois and Aubert 1985). Storm and 
Verts (1966) determined that the movements of a 
radio-tracked rabid Striped Skunk (Mephitis mephitis) 
in its last weeks of life were not statistically different 
from the movements of non-rabid skunks. Also, a 
radio-telemetry study by Artois and Aubert (1985) on 
three wild foxes inoculated with rabies indicated that 
these animals occupied a comparable home range 
before and during the phase at which the virus would 
have been shed. Rosatte (unpublished) found that 
during a trap-vaccinate-release study in eastern On- 
tario, movements of rabid Raccoons were not different 
from movements of non-rabid Raccoons. If the same 
rules hold true for Raccoons as they do for skunks or 
foxes, then rabid Raccoons are likely to encounter 
the same conspecifics as they would while they were 
healthy. This indicates that data obtained from this 
study have direct relevance to potential movements of 
Raccoons in the study population should they become 
infected with rabies. That is, dispersing yearling Rac- 
coons using the common feeding site would probably 
be the main vectors spreading the disease into areas 
beyond the population. 

Contact rates for Red Foxes (Vulpes vulpes) used 
in rabies simulation models are estimates of potential 
contact rates drawn from analysis of home range over- 
lap obtained from radio telemetry data (Blancou et 
al. 1991). The low precision of the telemetry system 
used in this study prevented analysis of home range 
overlap, spatial relationships and potential contact 
rates. It was not possible to tell, with the coarseness 
of the grid used for grid cell home range analysis, the 
potential for Raccoons with overlapping ranges to con- 
tact each other. Two Raccoons could be within the 
same 23-ha grid cell area at the same time and yet be 
unaware of each other. 

Data from this study indicated extensive overlap of 
home ranges within the population. Territoriality does 
not normally occur in Raccoons (Kauffmann 1982; 
Seidensticker et al. 1988) and has only been found for 
adult males in the spring and summer in North Dakota, 
at the northern edge of the Raccoons’ range and is 
thought to be due to competition for access to females 
(Fritzell 1978). Studies from more southern latitudes 
failed to demonstrate territoriality in Raccoons (Stuewer 
1943; Johnson 1970). 

Although caution should be used in comparing den- 
sities between studies, especially if different method- 
ologies, seasons, and habitats are involved, population 
densities for Raccoons are usually around one Rac- 
coon per 8-10 ha (Kauffmann 1982) but may range 
from one Raccoon per 100 ha in North Dakota and 
northern Ontario (Fritzell 1978; Rosatte 1996*) to one 


70 THE CANADIAN FIELD-NATURALIST 


Raccoon per ().4 ha in more favourable habitat (Twich- 
ell and Dill 1949). In southern Ontario, Raccoon 
density averaged over 200 plots sampled was one 
Raccoon/9-33 ha (Rosatte, unpublished) though den- 
sities of up to one Raccoon/1.8 ha have been recorded 
in forested park areas of Scarborough, Ontario (Rosatte 
et al. 1991). Density for our study area falls within the 
high end of the average range for Raccoon density in 
southern Ontario. It should also be noted that in our 
study, trapping for population estimates occurred at 
the end of summer when the local population ex- 
panded because the young-of-the-year were entering 
the population. 

Causes of mortality for Raccoons include starvation, 
heavy parasitism, poison, dogs, automobiles, hunting, 
trapping, canine distemper, and of course, rabies (Mech 
et al. 1968; Rosatte and MacInnes 1989; Riley et al. 
1998; Rosatte et al. 1991; Rosatte 2000); however, 
the hunting season in this area was from 15 October 
to 31 December (OMNR 1996*), after Raccoon den- 
sity was estimated in this study. Thus, trapping and 
hunting mortalities for the fall had yet to occur. Also, 
a considerable number of the dump Raccoons were 
trapped early in the study and vaccinated against rabies 
and distemper; therefore, the major source of mortal- 
ity in this population at the time of the study was 
probably due to collisions with automobiles. Apart 
from roadway mortality, the death rate of juveniles in 
this population was probably very low until winter 
food shortages set in. For this reason, the density 
estimated for our population probably represents a 
peak annual value. 


Management Implications 

Raccoon home range and movement data were used 
to estimate areas that need to be treated to control Rac- 
coon rabies in Ontario (Rosatte et al. 2001). Intimate 
knowledge of Raccoon behavior provides benchmarks 
to consider when determining the width of the popula- 
tion reduction and vaccination zones in order to pre- 
vent the spread of Raccoon rabies (Raccoon Rabies 
Task Force 1992*). These data are also being used to 
develop a Raccoon rabies model for Ontario which 
will be capable of predicting the movement of Raccoon 
rabies over time in the absence or presence of rabies 
control strategies. 


Acknowledgments 

We thank R. Harmsen, R. J. Robertson (Queen’s 
University), D. Voigt (Peterborough Ontario Ministry 
of Natural Resources: OMNR), and D. Johnston for 
helpful comments during the planning of the study and 
on early drafts of the manuscript. Field assistance was 
provided by L. Smail, J. Lugarich, and A. Thompson. 
Logistical support was provided by M. Power and D. 
Grieve, and Raccoons were aged by D. Joachim and 
A. Silver (Peterborough OMNR). Many thanks to A. 
Cross (Opinicon Resort) for allowing the researchers 
access to his property. And finally, thanks to F. Phelan 


Vol. 118 


and F. Connor at the Queen’s University Biological 
Station, and to M. Schunk (Queen’s University). This 
study was supported by an operating and equipment 
grant from the Ontario Ministry of Natural Resources. 
S. Totton received funding from an Ontario Graduate 
Scholarship and a Queen’s Graduate Award. 


Documents Cited (marked * in text) 
Ontario Ministry of Natural Resources. 1996. Hunting 
and Trapping Regulations for the 1996 hunting season. 
Raccoon Rabies Task Force. 1992. The raccoon strain of 
rabies: recommendations to prevent its becoming estab- 
lished in Ontario. Unpublished report. 25 pages. 

Rosatte, R. C. 1996. 1996 raccoon rabies research and con- 
trol annual report. Ontario Ministry of Natural Resources 
internal report. 


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Received 4 October 2001 
Accepted 20 May 2004 


Status of Marine Turtles in British Columbia Waters: A Reassessment 


DONALD F. MCALPINE! , STAN A. ORCHARD?, KELLY A. SENDALL?, and Rop PaLm* 


'New Brunswick Museum, 277 Douglas Avenue, Saint John, New Brunswick E2K 1E5 Canada dmcalpin @nb.aibn.com 
>3Royal British Columbia Museum, 675 Belleville Street, Victoria, British Columbia V8W 9W2 Canada 

‘Strawberry Isle Research Society, Box 213, Tofino, British Columbia VOR 2Z0 Canada 

>Present address: World Wide Fund for Nature, GPO Box 528, Sydney, NSW 2001 Australia 


McAlpine, Donald F., Stan A. Orchard, Kelly A. Sendall, and Rod Palm. 2004. Status of marine turtles in British Columbia 
waters: a reassessment. Canadian Field-Naturalist 118(1): 72-76. 


Marine turtles in British Columbia have previously been considered off course stragglers. Here we document 20 new reports 
for Green Turtles, Chelonia mydas, and Leatherback Turtles, Dermochelys coriacea, for the province. Until recently there 
had been no concerted effort to acquire data on marine turtle abundance or frequency off British Columbia. Observations 
presented here allow a reassessment of marine turtle status in British Columbia waters. We suggest Green Turtles and 
Leatherbacks should be considered rare vagrants and uncommon seasonal residents, respectively, off British Columbia and 
that they are a natural part of the British Columbia marine environment. 


Key Words: Green Turtle, Chelonia mydas, Leatherback Turtle, Dermochelys coriacea, British Columbia, status. 


There are few published reports for marine turtles 
in British Columbia waters. Marine turtles in the prov- 
ince have generally been considered “straggler[s]” 
(Kermode 1932) or “off course” (Gregory and Camp- 
bell 1984). Nussbaum et al. (1983) ignore marine tur- 
tles in their treatment of northwest amphibians and 
reptiles, and marine turtles are not included in a recent 
field guide to northwest reptiles “because sea turtles 
rarely visit these shores” (St. John 2002). During the 
preparation of a review of the status and conservation 
of marine turtles in Canadian waters (McAlpine et al. 
in press) recent occurrences of Green Turtles off Brit- 
ish Columbia were encountered (McAlpine et al. 2002). 
Here we document additional new reports for Green 
Turtles, Chelonia mydas, as well as Leatherback Tur- 
tles, Dermochelys coriacea, for British Columbia. These 
new records nearly double the number of published 
occurrences for marine turtles from the province. When 
considered in the context of a recent compilation for 
adjacent Alaska (Hodge and Wing 2000), these obser- 
vations suggest a reassessment of sea turtle status in 
British Columbia waters is warranted. The precipitous 
decline in Pacific Leatherback numbers (Spotila et al. 
2000), so serious it has led to a call for a moratorium 
on long-line fishing in the Pacific Ocean (Anonymous 
2002), also provides a timely context. 

Based on terms used to document bird status in 
Alaska, Wing and Hodge (2002) have proposed a 
standardized terminology to describe marine turtle 
occurrences. This terminology, ranging through seven 
categories from accidental to abundant, is loosely based 
on number of occurrences only. Although useful, this 
system ignores the ecological context for occurrence. 
Here we suggest modifying the system of Wing and 
Hodge (2002) by adding, where appropriate, one of 
four descriptors to their categories: resident, seasonal 


72 


resident, migrant, or vagrant. Resident refers to turtles 
present year round, seasonal resident refers to turtles 
present only at certain times of the year, migrant refers 
to turtles moving through a broad area to another loca- 
tion, vagrant refers to turtles occurring outside their 
normal resident or migratory range. We use this system, 
in conjunction with that of Wing and Hodge (2002), 
to describe marine turtle status in British Columbia. 
There are 11 published reports for Green Turtles 
(Carl 1955; Radovich 1961 cited in Stinson 1984; 
Hodge and Wing 2000; McAlpine et al. 2002) from 
British Columbia and 14 reports of Leatherbacks (Ker- 
mode 1932; Carl, 1944, 1960; MacAskie and Forester 
1962). Published and unpublished marine turtle reports 
for British Columbia prior to 1982 are summarized in 
Stinson (1984). Table | lists one unpublished British 
Columbia Green Turtle report, included by Stinson 
(1984) and six new ones; three unpublished 
Leatherback reports included in Stinson (1984) and 
10 new Leatherback occurrences. There are currently 
26 British Columbia reports for the Leatherback 
accompanied by sufficient locality data to map 
(McAlpine et al. in press), as well as other reports 
which are not site-specific. All of the 16 British Colum- 
bia Green Turtle reports can be map plotted (McAlpine 
et al. 2003). Several of the recent Green Turtle records 
are supported by photographic evidence (Figures 1- 
3), as are four of the Leatherback sightings (Figure 4; 
Table 1). The skull in Figure | is readily identified as 
that of C. mydas using the figures in Wyneken (2001). 
This stranding is reported in issue Number 2 of KSM 
Wutsiin, the official newletter of Hartley Bay School. 
Unfortunately, the skeletal remains of this turtle were 
not retained. Detailed necropsies carried out on the 
Green Point and Matlahaw Point C. mydas are now 
archived at the Royal British Columbia Provincial 


2004 


MCALPINE, ORCHARD, SENDALL, AND PALM: MARINE TURTLES 73 


TABLE |. Recent occurrences of marine turtles from British Columbia waters. 


Date Location 


Chelonia mydas 

22 November 1996 Hartley Bay 
early November 1998 mouth of Tlell R 
early November 1998 5 km S of Tlell R 
6 November 2001 4km N of Tlell R 
20 December 2001 Greene Point 

21 January 2002 Matlahaw Point 


Dermochelys coriacea 


mid-July 1970 Sharbau Island 
summer 1977 Off Ucluelet 
September 1977 Off Ucluelet 
30 August 1981 Skidiyate Inlet 


Hectate Strait 
Esperanza Inlet 


1 September 1981 
16 September 1982 


August 1993 Clerke Point 

26 May 1996 Mt Douglas Park 
May 1997 Kyuquot Sound 
September 1997 SE of Langara Island 
Spring 1998 Tofino 

6 September 2000 __ off shore 

4 August 2001 off Langara Island 


'PC = personal communication. 


3525) 
S336) 
SS nS9: 
SS esi) 
49° (3' 
49° 23' 


S130; 
49° 55' 
49° (0! 
Sul: 
53° 20' 
49° 52' 
50° 05' 
49° 08' 
50° 05' 
54° 12' 


49° 07' 
48° 43' 
54° 18' 


18" 


129215) 
131° 56' 
131° 54' 
ISH aps) 
125° 43' 
126° 29' 


128° 45' 
126° 38' 
125° 50’ 
131 730% 
130° 30' 
126° 44! 
127° 48' 
123° 58' 
W27e213\ 
132° 58' 


1252753: 
127° 26' 
1335210; 


Latitude(N) Longitude(W) Comments 


Carcass, Figure | 

Carcass, PC!: W. Flood to DFM 

Carcass, PC: W. Flood to DFM 

Fresh carcass, carapace 78.7 cm, Figure 2 
Fresh carcass, male, carapace 68.7 cm, Figure 3 
Carcass, male, carapace 69.9 cm 


Stinson (1984) 

Stinson (1984) 

Free swimming, RBCPM photo number 695 
Alive in salmon gillnet, RBCPM photo number 749 
Stinson (1984) 

RBCPM photo number 826 

Feeding on Cyanea, pc: J. Watson to G. Ellis 
Free swimming/Times-Colonist, 1 June, page B2 
Carcass, PC: G. Jamieson to DFM 

Free swimming, large numbers of Vellela sp. 
present, PC: S. Buchanan to S. Stebbins 
Decomposed carcass, PC: to SAO 

Free swimming, PC: K. Morgan to DFM 

Free swimming, PC: E. Simkin to G. Ellis, Figure 4 


Ficure 1. Skeletal remains of a dead Green Turtle stranded at Hartley Bay, British Columbia, 22 November 
1996. The skull in this photo can be readily identified as that of C. mydas (Hartley Bay School photo). 


74 THE CANADIAN FIELD-NATURALIST 


Museum and the Strawberry Isle Research Society, 
Tofino. A report prepared by a veterinary pathologist 
following examination of the Green Point Turtle states 
this turtle died from a bacterial infection to which it 
may have been predisposed following exposure to 
environmental stressors, such as low water tempera- 
ture. The animal was also suffering from a moderately 
severe case of pneumonia. The Matlahaw turtle ap- 
peared healthy and cause of death was not determined. 

Gregory and Campbell (1984) suggested the single 
live Green Turtle from British Columbia reported by 
Carl (1955) was “probably just off course”. On the basis 
of minimum thermal tolerance in cheloniid turtles and the 
few additional reports of live C. mydas. McAlpine et al. 
(2002) concurred, considering the species “accidental” 
in the province. Kermode (1932) identified the Leath- 
erback as a “‘straggler” in the North Pacific, Carl (1944) 
reported the species as “wandering” to British Colum- 
bia waters, and Cook (1981*) considered Leatherback 
Turtles observed in Canada were either “migrants or 
strays”. 

Stinson (1984), however, suggested occurrences of 
sea turtles in the northeastern Pacific could not be dis- 
missed as accidental but are influenced by temperature 
anomalies in the ocean. While she found sightings of 
marine turtles occur regularly during years of normal 
ocean temperatures, greater numbers of turtles were 
observed than expected when sea temperatures were 
above the mean. Hodge and Wing (2002) noted that 
in Alaska marine turtle occurrences are almost equally 
divided between warm-water and normal-water years. 
Most noteworthy was the complete lack of turtles in 
cold-water years. 

Whether the increased number of recent reports 
reflects a real increase in the prevalence of marine 
turtles off British Columbia, or simply increased public 
awareness and reporting, is unknown. However, based 
on their standardized occurrence terminology for 


< ESTOORG Ata oy AR RY a ARAN et os Ga 

FIGURE 2. This 78.7 cm Green Turtle came ashore fresh 
dead on 6 November 2001, 4 km N of the Tlell 
River, Queen Charlotte Islands, British Columbia 
(V. Flood photo). 


Vol. 118 


FIGURE 3. This 68.7 cm male Green Turtle came ashore fresh 
dead at Green Point, Vancouver Island, British Col- 
umbia, on 20 December 2001. (P. Clarkson/Parks Can- 
ada photo). 


marine turtles, Wing and Hodge (2002) report that 
marine turtles occur too often in Alaska waters to be 
considered accidental. These authors consider marine 
turtles a natural part of the Alaskan marine environ- 
ment. Likewise, Stinson (1984) concludes marine 
turtles are a regular part of the environment in the 
northeast Pacific. Based on the frequency of reports, 
Wing and Hodge (2002) considered the Green Turtle 
and Leatherback Turtle as rare and uncommon, respec- 
tively, in Alaska. Stinson (1984) identified a marine 
turtle season of July to September in the northeast 
Pacific Ocean and stated September was the critical 
marine turtle month in British Columbia waters. How- 
ever, nearly all of her British Columbia marine turtle 
observations are of Leatherbacks. Currently, the sea- 
sonal pattern of occurrence that appears to be emerg- 
ing, both in British Columbia and Alaska, suggests 
that D. coriacea is indeed most frequent during the 
July to September period, while C. mydas is most 
likely to be found stranded in the northeast Pacific 
during October to December. It should be noted that 
most British Columbia Green Turtle observations have 
been of stranded turtles, often recently deceased. 
Hodge and Wing (2000) observe that the low 
frequency of hard shell turtles from Alaska waters 
indicates these turtles are straying beyond their toler- 
able range, while they consider Dermochelys in Alaska 
as occupying marginal habitat that may not be con- 
sistently used from year to year. The accumulating 
number of marine turtle observations from British 
Columbia suggests that these species are even more 
frequent in British Columbia than Alaska. Following 


2004 


MCALPINE, ORCHARD, SENDALL, AND PALM: MARINE TURTLES 75 


Ficure 4. Leatherback turtle photographed 4 August 2001 off Langara Island, Queen Charlotte Islands, British Columbia 
(Erik Simkin photo). 


Wing and Hodge (2002), and our own suggestions 
above, we describe C. mydas as a rare vagrant and 
and D. coriacea as an uncommon seasonal resident 
in British Columbia waters. However, we would add 
there has been no effort to methodically collect infor- 
mation from dead stranded turtles nor, until a recent 
initiative by the Vancouver Aquarium (C. Sbrocchi, 
personal communication to DFM), to acquire sightings 
data on marine turtle abundance or frequency off 
British Columbia. Anecdotal information on several 
Leatherback occurrences is included in the KSM 
Wutsiin newsletter noted above, indicating canvas- 
sing local communities, and especially native fishers, 
could be a source of information on marine turtle 
occurrences in British Columbia. Additionally, Glen 
Jamieson (Pacific Biological Station, personal commu- 
nication to DFM) reports that Leatherbacks are 
“periodically seen off the west coast of [Vancouver 
Island]” and Frank Bernard (Pacific Biological Station, 
personal communication to SAO) reported that he had 
seen as many as six Leatherbacks in a day 80-320 km 
off the west coast of Vancouver Island. Reports of the 
Pacific Ridley, Lepidochelys olivacea, and the Logger- 
head, Caretta caretta, for Alaska (Hodge and Wing 
2000) suggest focused investigations may reveal these 
species also occur off the British Columbia coast. 


Acknowledgments 

We thank the following individuals for sharing 
British Columbia marine turtle observations and infor- 
mation with us and allowing us to include their reports 
here: Scott Buchanan, Barry Campbell, Graeme Ellis, 
Glen Jamieson, Marilyn Joyce, Wayne and Virginia 
Flood, Ken Morgan, Carla Sbrocchi, Eric Simkin, Jane 
Watson, Kris Willcock, and the late Frank Bernard. 
In particular, we would like to thank Eric Hill and 
Simone Westgarth of the Hartley Bay School for their 
efforts in obtaining information that allowed identifi- 
cation of the Hartley Bay turtle. 


Documents Cited (marked * in text) 

Cook F. R. 1981. Status report on the Leatherback Turtle, 
Dermochelys coriacea. Committee on the Status of En- 
dangered Wildlife in Canada, Ottawa, 17 pages. 


Literature Cited 

Anonymous. 2002. Moratorium on Pacific long-lining could 
save leatherbacks. Marine Turtle Newsletter 97: 26. 

Carl, G. C. 1944. The reptiles of British Columbia. Hand- 
book Number 3. British Columbia Provincial Museum, 
Victoria, British Columbia. 

Carl, G. C. 1955. The green turtle in British Columbia. 
Report of the Provincial Museum of Natural History and 
Anthropology, Victoria, British Columbia 1954: B77-78. 


76 THE CANADIAN FIELD-NATURALIST 


Carl, G. C. 1960. The reptiles of British Columbia. Hand- 
book Number 3. British Columbia Provincial Museum, 
Victoria, British Columbia. 

Gregory, P. T., and R. W. Campbell. 1984. The reptiles of 
British Columbia. Handbook number 44. British Colum- 
bia Provincial Museum, Victoria, British Columbia. 

Hodge, R. P., and B. L. Wing. 2000. Occurrences of marine 
turtles in Alaska waters 1960-1998. Herpetological Review 
31: 148-151. 

Kermode, F. 1932. A remarkable capture of Leatherback 
Turtles off Bajo Reef, near Nootka Sound, West Coast of 
Vancouver Island, British Columbia. Report of the Prov- 
incial Museum of Natural History for the year 1931. 
Victoria, B.C. Pages 6-7. 

MacAskie, I. B., and C. R. Forrester. 1962. Pacific Leather- 
back off the coast of British Columbia. Copeia 1962: 646. 

McAlpine, D. F., M. C. James, J. Lien, and S. A. Orchard. 
In press, Status and conservation of marine turtles in 
Canadian waters. Jn Ecology, conservation and status of 
reptiles in Canada. Edited by C. R. Seburn and C. A. 
Bishop. Herpetological Conservation 3. 

McAlpine, D. F., S. A. Orchard, and K. A. Sendall. 2002. 
Recent occurrences of the green turtle from British Colum- 
bia waters. Northwest Science 76: 185-188. 


Vol. 118 


Nussbaum, R. A., E. D. Brodie Jr., and R. M. Storm. 
1983. Amphibians and reptiles of the Pacific Northwest. 
University of Idaho Press, Moscow, Idaho. 

Radovich, J. 1961. Relationship of some marine organisms 
of the northeast Pacific to water temperatures, particularly 
during 1957 through 1959. California Fish and Game, 
Fisheries Bulletin 112: 1-62. 

Spotila, J. R., R. D. Reina, A. C. Steyermark, P. T. Plotkin, 
and F. V. Paladino. 2000. Pacific Leatherback Turtles face 
extinction. Nature 405: 529-530. 

Stinson, M. L. 1984. Biology of sea turtles in San Diego Bay, 
California and in the northeastern Pacific Ocean. MSc. 
thesis, San Diego State University, California. 

St. John, A. 2002. Reptiles of the Northwest: British Colum- 
bia to California. Lone Pine Publishing, Edmonton, Alberta. 

Wing, B. L., and R. P. Hodge. 2002. Occurrence terminology 
for marine turtles. Marine Turtle Newsletter 95: 15-16. 

Wyneken, J. 2001. The anatomy of sea turtles. U.S. Depart- 
ment of Commerce, NOAA Technical Memorandum 
NMEFS-SEFSC-470. 172 pages. 


Received 27 February 2003 
Accepted 20 November 2003 


| 


| 


Introduced Marine Species in the Haida Gwaii (Queen Charlotte Islands) 
Region, British Columbia 


N. A. SLOAN and P. M. BARTIER 


Parks Canada, Gwaii Haanas National Park Reserve and Haida Heritage Site, P.O. Box 37, Queen Charlotte, British Columbia, 
VOT 1S0 Canada 


Sloan, N. A., and P. M. Bartier. 2004. Introduced marine species in the Haida Gwaii (Queen Charlotte Islands) region, British 
Columbia. Canadian Field-Naturalist 118(1): 77-84. 


This historical review of a marine area’s introduced species was facilitated by geo-referenced marine species inventories of the 
Haida Gwaii (Queen Charlotte Islands) region. One plant, 14 invertebrate, and two fish introduced species have been recorded 
since the early 20" century from the marine waters around Haida Gwaii. Records of species occurrences are listed and mapped, 
and modes of introduction are discussed. It will be important to continue documenting areas’ introduced species locations to 


track the progress of invasions that could affect local marine ecosystem well-being. 


Key Words: introduced species, marine, Haida Gwati, Queen Charlotte Islands, British Columbia. 


*“... the control of alien marine species is in its infancy.” 


(Bax et al. 2001) 


Introduced (non-indigenous) marine species are of 
global concern (Bax et al. 2001). The dynamism and 
connectivity of marine ecosystems, mariculture and 
shipping facilitate species’ introductions. The introduc- 
tion of some species may be followed by rapid local 
dispersal of propagules and appreciable ecological con- 
sequences (Grosholz 2002). Introduced species know]- 
edge is more developed for land and freshwater than 
for marine ecosystems (Ruiz et al. 2000; Simberloff 
2000; Bax et al. 2001). 

Carl and Guiguet (1958) were the first to broadly 
document species introductions in Pacific Canada. Cur- 
rent awareness of the problem is growing in the region, 
and Levings et al. (2002) have reviewed introduced 
marine species found in the Strait of Georgia, south- 
ern British Columbia. 

Using the 30 introduced mollusks along the Pacific 
coast of North America as an example, Carlton (1992) 
reported the main modes of introduction as: (1) asso- 
ciated with introduced Japanese and Atlantic oysters 
imported for mariculture (approximately 27 species), 
(2) via vessel hull fouling and organisms living in wood, 
and (3) from overseas ships’ ballast water releases. 
More recently, Chapman et al. (2003) reported that 
along the U.S. Pacific coast, 24 of the 37 marine and 
estuarine bivalve species commercially available as sea- 
food are alien species. 

We report on the documented presence of introduced 
marine plants, invertebrates and fishes in the Haida 
Gwatii (Queen Charlotte Islands) region of northern 
British Columbia. No introduced marine birds or mam- 
mals have been recorded. The plant and invertebrate 
records, gleaned from the literature and museum col- 
lections, are archived in the geographic information 
system (GIS) databases of Gwaii Haanas National Park 
Reserve and Haida Heritage Site (Sloan and Bartier 


2000; Sloan et al. 2001). We excluded introduced 
species whose total geographic range encompasses the 
Haida Gwaii region, although, of course, their presence 
could be inferred. An example is the amphipod Coro- 
phium acherusicum from Asia and now known from the 
northern mainland British Columbia coast (Bousfield 
and Hoover 1997). The American Shad (Alosa sapidis- 
sima) have long been known from the whole northeast 
Pacific (Welander 1940), but only recently from Haida 
Gwaii waters (Workman et al. 1996). Atlantic salmon 
(Salmo salar) observations were from the Atlantic sal- 
mon Watch Program web site: http://www.pac.dfo-mpo. 
gc.ca/sci/aqua/AS WP_e.htm. The fish records are not 
yet in our database. 


Results and Discussion 

For Haida Gwaii, accidental (or active) introductions 
of one plant, 14 invertebrate, and two fish species asso- 
ciated with fishery or mariculture development are 
summarized in Table 1 and illustrated in Figures | and 
2. Likely some of these species have spread northward 
to Haida Gwaii from the more developed southern 
mainland British Columbia and U.S. coasts where they 
first became established. There have been introductions 
of species to British Columbia targeted for mariculture, 
such as Pacific Oyster (Crassostrea gigas) from Japan, 
plus their attached associates (Quayle 1988) or parasites 
(Bower et al. 1994). For example, the seaweed Sargas- 
sum muticum is now ubiquitous coast-wide, including 
Haida Gwaii (Figure 1). It was accidentally introduced 
into southern British Columbia in the early 20" century 
attached to Pacific oysters. Another example of collateral 
introduction in British Columbia is the parasitic copepod 
Mytilicola orientalis, likely introduced via Pacific Oyster 
stock and now widely infesting Native Littleneck Clams 
(Protothaca staminea) and Butter Clams (Saxidomus 
giganteus) in southern British Columbia (Bower et al. 
1994). 


TL 


Vol. 118 


THE CANADIAN FIELD-NATURALIST 


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79 


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: INTRODUCED MARINE 


SLOAN AND BARTIER 


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80 THE CANADIAN FIELD-NATURALIST 


Langara Island 
Parry Passage 
Cryplosula pallasiana 
Lininodrilus monothecus 


Cryptosula pallasiana 


Graham 


Rennell Sound Island 


Rose 


Masset Spit 
Delkatla 


Vol. 118 


° Pelonaia corrugata 


Trechus oblusus 


Limnodrilus nonothecus 


Kumdis Bay 


Yakoun Bay 


a 


Skidegate 
Inlet 


Moresby 
Island 


Cartwright 
Sound 


Tasu 
Sound 


Bowerbankia gracilis 
Sabia conica 


Ciona intestinalis 


Legend 


a Sargassum mulicum 
Mya arenaria 
Ampithoe vallida 


® Other invertebrate sightings as noted 


FiGure |. Map of Haida Gwaii showing locations mentioned in the text and collection sites of introduced marine plant and 
invertebrate species, based on data from Sloan and Bartier (2000) and Sloan et al. (2001). 


Mariculture-associated Introductions 


In 1977, concern over introduced aquatic species 
issues stimulated establishment of the federal-prov- 
incial Fish Transplant Committee (recently renamed 
Introductions and Transfers Committee — ITC) under 


Bowerbankia gracilis _ 


Schizoporella unicornis ® 


Tubularia crocea 


mandates from the federal Fisheries Act and the Bri- 
tish Columbia Fisheries Act and British Columbia Wild- 
life Act (BC 1990). The ITC evaluates potential risks 
to the environment associated with introductions or 
transfers of either finfish or invertebrates (“shellfish’’) 


Hecate 


Strait 


Hotspring 
Island 


Dolomite "Burnaby" 
Narrows 


Harriet 
Harbour 


Houston Stewart 
Channel 


2004 SLOAN AND BARTIER: INTRODUCED MARINE SPECIES 8] 


4 row 
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FiGurE 2. Map of the northern British Columbia and southeast Alaska regions showing the density of Atlantic salmon obser- 
vations. All data are from Fisheries and Oceans Canada’s Atlantic Salmon Watch Program: http://www.pac.dfo-mpo.gc. 
ca/sci/aqua/ASWP_e.htm. [accessed May, 2004]. British Columbia data are from 1987 to 2002 and partitioned according 
to Pacific Fishery Management Areas, and the Alaska data are from 1990 to 2002 and partitioned according to the 
Alaska Department of Fish and Game Area Polygons. 


into British Columbia marine or freshwaters. Besides _ genetic material into native species. Further, there are 
the species themselves and their attached associates, concerns over potential ecological displacement of 
there is the risk of introducing diseases, parasites or native species. The ITC issues licences to introduce 


82 THE CANADIAN FIELD-NATURALIST 


aquatic species into British Columbia or to transfer 
species between domestic water bodies. 

Shellfish mariculture remains a potential avenue of 
species introduction to Haida Gwaii. A Masset-based 
group has investigated the potential for local shellfish 
mariculture for coastal community economic develop- 
ment. Leased culture operations in Skidegate Inlet and 
Rennell Sound remain active for Pacific Oyster culture. 
These operations are unlikely a threat of introducing 
oysters as local waters are too cold for oyster breeding 
although they are suitable for growth of certified 
disease-free juveniles (spat) from culture in southern 
British Columbia. However, local waters may not be 
too cold for oysters’ parasites or other associated 
species. From 1997 to 2001, pilot raft culture sites 
were tested to grow certified disease-free Weathervane 
Scallop (Patinopecten sp.) hybrid (native x Japanese) 
spat (B. Mark, Masset, personal communication). Only 
the Rennell Sound site remains active for scallop cul- 
ture (R. Lozon, Queen Charlotte City, personal com- 
munication). The potential for species introduction is 
likely low, because no successful settlement of hybrid 
Weathervane Scallops has occurred in British Colum- 
bia in the last decade (Island Scallops Ltd., Qualicum 
Beach, personal communication). 


Ships’ Ballast Water 


Gauthier and Steel (1998) reported that Canada was 
receiving approximately 52 million tonnes of ballast 
water from foreign shipping annually with little pro- 
tective policy or regulation. Indeed, the major vector 
of introductions to Pacific North America has been 
from shipping (Ruiz et al. 2000). Concerning Pacific 
Canada, Levings (1999) mentioned the Canadian Bal- 
last Water Management Guidelines issued by Transport 
Canada in 2000 with an Annex (II) for the Pacific 
coast aimed at preventing introduction of non-indige- 
nous aquatic organisms (http://www.tc.gc.ca/Marine 
Safety/). These Guidelines will become Regulations 
under the Canada Shipping Act. 

The Vancouver Port Authority has had a mandatory 
ballast water program since 1997. It is based on the 
assumption that mid-ocean ballast water exchange, with 
water containing pelagic species not likely adapted to 
coastal conditions, decreases likelihood of introducing 
viable species into port waters (Levings et al. 2004). 
Port authorities are now finding, however, that such ex- 
change criteria are only partially effective (C. Levings, 
Fisheries and Oceans Canada (DFO), personal com- 
munication). Other British Columbia ports invoking 
ballast water management are Nanaimo and New West- 
minster. The north coast of British Columbia, with an 
active deep-water international port in Prince Rupert, 
however, has no ballast water program. Further, Lev- 
ings et al. (2004) express concern about “intracoas- 
tal” transport. For example, from Haida Gwaii there is 
on-going barge and self-dumping log barge traffic with 


Vol. 118 


southern British Columbia and there has been relative- 
ly recent international vessel traffic for mine ore con- 
centrates from Tasu Sound (mine closed 1973) and 
Harriet Harbour (mine closed 1968). 

The ecological effects of introduced species such as 
invertebrates on the Pacific coast are poorly studied 
(Carlton 1992). It is sobering to reflect that, once in- 
troduced, marine species may be difficult to control 
and their ecosystem consequences may be damaging 
(Simberloff 2000; Grosholz 2002). An example is the 
European Green Crab, Carcinus maenas, introduced 
to the San Francisco Bay area in 1989. The Green Crab 
has since been recorded from Esperanza Inlet on the 
northwest coast of Vancouver Island (Jamieson et al. 
2002). As active predators, Green Crabs could affect 
British Columbia intertidal fauna, as they have in 
California (Grosholz 2000). 

Times and attitudes have changed about introduc- 
tions of marine species. In the 1980s, DFO discussed 
introduction of the large, predatory Atlantic lobster 
(Homarus americanus) around Haida Gwati. Barber 
(1983) concluded that up to 7700 km? of Hecate Strait 
area south of Skidegate Inlet, as well as Masset Inlet, 
were suitable for lobster introduction. Now, such ini- 
tiatives are contrary to agency mandates for sustain- 
able, ecosystem-based management. 

Among introduced marine species, Atlantic Salmon 
(Salmo salar) currently has the highest ecosystem 
and socio-political profile in British Columbia (Gross 
1998; Volpe et al. 2001). Important issues include 
escapement and persistence in river systems facilitating 
competition for spawning habitat with native salmo- 
nids. Further, the threat of continuing introductions 
helps animate the vigorous debate over expansion of 
salmon farming in British Columbia. Although Atlantic 
Salmon have been reported from Haida Gwaii coastal 
waters, there are no salmon farms in Haida Gwaii, and 
there are no records at this time of Atlantic salmon from 
within Haida Gwaii rivers and streams (V. Fradette, 
DFO, personal communication). 

This overview of a marine region’s introduced spe- 
cies was facilitated through marine species inventories 
from the historical literature of the Haida Gwaii region. 
We do not claim that it is complete, but it is a start. It 
will be important to continue documenting introduced 
species reports to track the progress of introductions 
that could affect local marine ecosystem well-being. 


Acknowledgments 

We thank C. D. Levings of Fisheries and Oceans 
Canada (DFO) for reviewing an early draft and G. E. 
Gillespie (DFO) for information on fishes. 


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Received 27 December 2002 
Accepted 11 June 2004 


Ruby-throated Hummingbird, Archilochus colubris, Entanglements in 
Burdock, Arctium spp., at Delta Marsh, Manitoba 


HEATHER L. HINAM!, SPENCER G. SEALY, AND Topp J. UNDERWOOD 


Department of Zoology, University of Manitoba, Winnipeg, Manitoba R3T 2N2 Canada 
'Present Address: Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9 Canada 
Corresponding author sgsealy @cc.umanitoba.ca 


Hinam, Heather L., Spencer G. Sealy, and Todd J. Underwood. 2004. Ruby-throated Hummingbird, Archilochus colubris, 
entanglements in burdock, Arctium spp., at Delta Marsh, Manitoba. Canadian Field-Naturalist 118(1): 85-89. 


Exotic burdock (Arctium spp.) pose a risk of mortality for small native birds, such as the Ruby-throated Hummingbird (Archi- 
lochus colubris), which may become entangled in its burrs. At Delta Marsh, Manitoba, we found 11 hummingbirds and five 
individuals of four species of songbirds entangled on burdock in the dune-ridge forest over a 20-year period. Entangled birds 
were mostly migrants. Most hummingbirds caught were juvenile males, whereas the few songbirds were mostly adult males. 
We suspect that hummingbird entanglements resulted from an attraction to the purple flowers of burdock, but aggressive 
interactions with conspecifics and other factors may have been involved. Birds may be at a higher risk of entanglement at 
important migratory stopover sites, such as Delta Marsh, where both burdock and large numbers of birds are concentrated in 
a small area. 


Key Words: exotic plants, burdock, Arctium spp., fatal entanglement, Ruby-throated Hummingbird, Archilochus colubris, pas- 


serines, mortality, Delta Marsh, Manitoba. 


Exotic species often exert negative effects on the 
survival of native flora and fauna (Atkinson 1989). This 
is exemplified by the relationship between burdock 
(Arctium spp.) and small birds and bats in North Amer- 
ica (McNicholl 1988, 1994). Four species of burdock 
were introduced from Eurasia into North America in 
the early 1600s (Gross et al. 1980; Harms 2001). All 
four species of burdock develop seed heads covered 
with burrs that stick to passing animals, thus dis- 
persing the seeds (McNicholl 1988). These burrs are 
hazards for small birds and bats because individuals 
may become entangled in the hooked bracts and die 
(McNicholl 1988, 1994). The effect of such entangle- 
ments on bird populations may be magnified when 
the plants are concentrated in small areas where birds 
nest or forage during migration. 

For many songbirds, the dune-ridge forest that sepa- 
rates the south shore of Lake Manitoba from Delta 
Marsh (50°11 N, 98°19’ W), Manitoba, is an important 
breeding area (Goossen and Sealy 1982; MacKenzie 
et al. 1982) and migratory stopover site (den Haan 
1996). Three species of burdock (A. minus, A. lappa, 
A. tomentosum) have been documented in the area, but 
Common Burdock (A. minus) is the most abundant, 
occurring along the edge of the marsh and in the ridge 
forest, as well as in ditches and at other disturbed 
sites (Shay 1999). The composition of the understory 
of the ridge forest has changed over the last 30 years 
and the amount of burdock apparently has increased 
(Kenkel and Graham 1994; S.G. Sealy, personal obser- 
vations). With this apparent increase, Delta Marsh has 
the potential to become a trap line for small birds such 
as the Ruby-throated Hummingbird (Archilochus colu- 
bris). Indeed, this species has been one of the most 


frequently discovered entangled in burdock (McNicholl 
1988, 1994; Raloff 1998; Nealen and Nealen 2000). 
Here we document additional cases of mortality of 
Ruby-throated Hummingbirds and songbirds due to 
entrapment on the seed heads of burdock at Delta 
Marsh, Manitoba. 


Entanglements 

Sealy and co-workers have studied songbird popu- 
lations in the ridge forest at Delta Marsh since 1973, 
but it was not until 1983 that the first bird was found 
entangled on burdock. The decomposed remains of a 
Ruby-throated Hummingbird were discovered that 
spring, but death had occurred the previous year. Since 
then, 10 more hummingbirds have been found, all but 
one in August 1985 (Table 1). One of these was alive 
and released, leaving behind several feathers stuck to 
the burrs (Table 1). From 1977 to 1979, 1981 and 1983, 
hummingbirds were banded in conjunction with gen- 
eral songbird banding in the ridge forest from mid-May 
through the end of August. The timing of the entan- 
glements in 1985 coincided with the fall migration of 
hummingbirds through the ridge forest in August 
(Figurel). However, except for the live hummingbird, 
the dates of entanglement only approximate the time of 
death because the rates of decomposition and mummi- 
fication are not known. Thus, some hummingbirds 
could have been killed earlier in August. 

Despite sporadic searching since 1985, only one 
other entangled hummingbird was found, in 2002 
(Table 1, Figure 2), but since 1994, five individuals of 
four other species have been found entangled fatally 
(Table 2). Another record consisted of only a few 
feathers from an unknown passerine species, possibly 


85 


86 THE CANADIAN FIELD-NATURALIST Vol. 118 


TABLE |. Records of Ruby-throated Hummingbirds entangled in burdock at Delta Marsh, Manitoba. 


Date Age/Sex! Condition Specimen No.” 
19 May 1983 Unk/unk Decomposed: overwintered, wings, ventral feathers attached 4986 
15 August 1985 Unk/unk Released, feathers stuck on burdock 29817 
18 August 1985 HY/female Recently caught (intact) 4982 
18 August 1985 HY/male Recently caught (intact) 4983 
18 August 1985 Unk/unk Partially decomposed: attached by ventral and wing feathers 4985 
19 August 1985 HY/female Recently caught 4981 
19 August 1985 AHY/male Partly decomposed 4984 
19 August 1985 AHY/male, — Facing each other with burrs in between 
HY/male (fresh and mostly intact) 4987 A,B 

21 August 1985 HY/male Stuck across burr (fresh, mostly intact) 4980 
14 September 2002 HY/female Decomposed, sternum visible 499] 


"Unk = unknown, HY = hatch year, AHY = after hatch year. 
Bird specimens deposited in the vertebrate collections of the Manitoba Museum, Winnipeg, Manitoba, Canada. 
>Feathers deposited in the University of Manitoba Zoology Museum, Winnipeg, Manitoba, Canada. 


TABLE 2. Records of songbird species entangled in burdock at Delta Marsh, Manitoba. 


Species Date Age/Sex! Condition Specimen No.” 
Unknown passerine 15 August 1985 Unk/unk Feathers stuck on burr 29823 
Golden-crowned Kinglet 30 October 1994 Unk/male Wings and breast attached (fairly intact) 4976 
Ruby-crowned Kinglet 2 November 1996 AHY/male Left wing caught (fresh, intact) 4977 
Common Yellowthroat May 1999 AHY/female | Decomposed: overwintered, 

caught by one leg 4978 

13 May 2000 AHY/male Decomposed: overwintered, caught 

by breast, both wings 4979 
Yellow-rumped Warbler? —_10 October 2002 HY/unk Fresh, caught by right foot, feathers 

of right flank, left wing 4988 


'Unk = unknown, HY = hatch year, AHY = after hatch year. 


*Bird specimens deposited in the vertebrate collections of the Manitoba Museum, Winnipeg, Manitoba, Canada. 
Feathers deposited in the University of Manitoba Zoology Museum, Winnipeg, Manitoba, Canada. 


‘Banded 14 September 2002. 


a sparrow (C. Dove, personal communication). This 
bird may have become entangled and freed itself; alter- 
natively it may have been preyed upon or scavenged. 
Two Common Yellowthroats (Geothlypis trichas), a 
species that nests at Delta Marsh, were found in spring 
after apparently becoming entangled the previous year. 
One Golden-crowned Kinglet (Regulus satrapa) and 
one Ruby-crowned Kinglet (R. calendula), both boreal 
forest breeders and late migrants, were found freshly 
dead in the fall. In October 2002, a freshly dead 
Yellow-rumped Warbler (Dendroica coronata) was dis- 
covered less than one month after it had been banded 
in the ridge forest. All four of these species have been 
found trapped on burdock elsewhere (McNicholl 
1988, 1994). 

The sex and age of birds were identified where pos- 
sible by characteristics of the plumage, feathers and 
bills (Pyle 1997). Of the hummingbirds entangled, five 
were males and three were females, whereas three 
songbirds were males and one was a female (Tables 
1, 2). Six hummingbirds were juveniles and two were 
adults (Table 1). By contrast, three songbirds were 
adults and only one was a juvenile (Table 2). 


Discussion 

Most birds entangled in burdock at Delta Marsh 
likely were migrants. Although Ruby-throated Hum- 
mingbirds nest at Delta Marsh (Underwood and den 
Haan 2000) in small numbers, all entanglements were 
discovered during their fall migration period through 
Delta Marsh (Figure 1). Among the other species killed 
on burdock, only the Common Yellowthroat breeds 
commonly at Delta Marsh (Underwood and den Haan 
2000). The preponderance of migrants in our sample 
of burdock mortalities is interesting. Several small 
songbirds, such as the Least Flycather (Empidonax 
minimus) and Yellow Warbler (Dendroica petechia), 
nest at high densities in the ridge forest (Goossen and 
Sealy 1982; Briskie and Sealy 1989; S. G. Sealy, un- 
published data), but neither species has been recorded 
entangled in burdock there, although the Least Fly- 
catcher is a documented victim elsewhere (Underwood 
and Underwood 2001). Individuals of both species have 
been observed perched on stems of burdock (Sealy and 
Underwood, personal observations) and Sealy ob- 
served two male Yellow Warblers, one chasing the other 


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HINAM, SEALY, AND UNDERWOOD: HUMMINGBIRD ENTANGLEMENTS IN BURDOCK 


87 


13 16 19 22 


August 


FicureE 1. Number of Ruby-throated Hummingbirds found caught on burdock at Delta Marsh, August 1985 (1 = 9), in 
relation to the number of hummingbirds banded per day throughout 1977 (n = 7), 1978 (n = 37) and 1979 (n= 4), 


1981 (n = 23) and 1982 (n = 42). 


that hit seed heads during the chase. Although they 
became stuck, they extricated themselves within sec- 
onds of impact. This observation notwithstanding, 
accidental strikes on burdock seed heads likely account 
for few mortalities. This may explain the infrequency 
of entanglements among the resident songbirds, which 
have not been observed foraging on burdock flowers 
or arthropods attracted to them. 

It is commonly assumed that birds become entangled 
while foraging (McNicholl 1988). Although humming- 
birds feed mainly while hovering (Robinson et al. 
1996), their small size may be the reason they frequent- 
ly become entangled. As noted by Nealen and Nealen 
(2000), Ruby-throated Hummingbirds prefer reddish 
flowers (Robinson et al. 1996; but see Miller and Miller 
1971), although they do not specialize on a particular 
species (Bertin 1982). Because burdock flowers are with- 
in the red spectrum (Gross et al. 1980), hummingbirds 
may be attracted to them in the ridge forest where there 
are few of the hummingbird’s putatively preferred spe- 
cies (Robinson et al. 1996; Shay 1999). 

Birds may also become entangled while foraging 
for insects that inhabit the flowers or the seed heads. 
Insects may comprise up to 60% of the diet of Ruby- 


throated Hummingbirds and individuals have been 
known to glean larval lepidoptera and other insects 
from the surface of plants (Robinson et al. 1996). Over 
20 species of insects from three Orders, Lepidoptera, 
Coleoptera and Hymenoptera, have been found on 
burdock seed heads (Mulligan and Kevan 1973; Gross 
et al. 1980). The kinglets and warblers may have been 
attempting to take insects from the burrs when they 
became entangled (see Needham 1909). 

Entanglements may also result from social inter- 
actions between individuals. The deaths of two male 
hummingbirds on the same burr cluster (Table 1, MM 
4987A and B) may have resulted when one individ- 
ual attacked the other at the flower cluster and they both 
became entangled as the interaction ensued. Sealy 
watched a male Ruby-throated Hummingbird attack 
another male as it hovered by a seed head. The hov- 
ering bird became entangled by one leg, but extri- 
cated itself within seconds. The attacker had already 
flown away. Wind may also cause entanglements, buf- 
feting birds against seed heads as they move through 
the area (McNicholl 1988). Thus, identifying the factor 
or combination of factors that promote individual bird 
entanglements is difficult. 


88 THE CANADIAN FIELD-NATURALIST 


FIGURE 2. Ruby-throated Hummingbird entangled in 
burdock found 14 September 2002, Delta Marsh 
(photograph by T. J. Underwood). 


The large number of hummingbird entanglements 
in burdock over a short period in August 1985 seems 
unusual. Burdock apparently has increased in abun- 
dance on the ridge forest (Kenkel and Graham 1994) 
since our studies began at Delta in 1973. This suggests 
that the frequency of entanglement should have in- 
creased or at least remained constant. However, search 
effort has varied widely since 1985 and migration 
rates and weather patterns affecting bird movements 
through the ridge forest vary from year to year. Hence, 
it is difficult to assess from occasional records the 
overall effect of burdock at Delta Marsh on migrating 
birds. Nevertheless, the growing number of reports of 
burdock-related deaths in birds (e.g., McNicholl 1994; 
Raloff 1998; Underwood and Underwood 2001) sug- 
gests that this type of mortality may be more important 
than originally believed, particularly at places like Delta 
Marsh, and King’s Park in Winnipeg (Underwood and 
Underwood 2001), where burdock and large numbers 
of migrating birds are concentrated. Further study of 
the interaction between birds and burdock and the pos- 
sibly fatal consequences for birds should result in a 
better understanding of the effects of this exotic plant 
species on bird populations. 


Acknowledgments 
We thank Janis Klapecki for cataloguing bird speci- 
mens in The Manitoba Museum and Carla Dove, 


Vol. 118 


Smithsonian Institution, for attempting to identify 
feathers. Some specimens were collected by G. C. 
Biermann, J. V. Briskie, H. E. den Hann, K. A. Hobson, 
E. Kiviat, A. Mcllraith, and R. M. Underwood. H. E. 
den Haan provided details on the banding of the Yel- 
low-rumped Warbler from the files of the Delta Marsh 
Bird Observatory and assisted with age/sex identifica- 
tion of specimens. We thank the reviews, A. J. Erskine 
and M. K. McNicholl, for their comments on the man- 
uscript. Accommodation and facilities were provided 
by the Delta Marsh Field Station (University of Mani- 
toba). Officers of the Portage Country Club permitted 
us to conduct some of the field work on their property. 
Funding was provided by the Natural Sciences and 
Engineering Research Council of Canada. 


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Miller, R. S., and R. E. Miller. 1971. Feeding activity and 
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Needham, J. G. 1909. Kinglets captured by burdocks. Bird- 
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Shay, J. M. 1999. Annotated vascular plant species list for 
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HINAM, SEALY, AND UNDERWOOD: HUMMINGBIRD ENTANGLEMENTS IN BURDOCK 89 


sity of Manitoba Field Station (Delta Marsh), Occasional 
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Underwood, T. J., and H. E. den Haan. 2000. Checklist of 
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Received 7 March 2003 
Accepted 23 April 2004 


Multiple Mating Results in Multiple Paternity in Richardson’s Ground 
Squirrels, Spermophilus richardsonii 


James F. Hare!, GLENDA Topp!, and WENDY A. UNTEREINER? 


' Department of Zoology, University of Manitoba, Winnipeg, Manitoba R3T 2N2 Canada 
* Botany Department, Brandon University, Brandon, Manitoba R7A 6A9 Canada 


Hare, James F., Glenda Todd, and Wendy A. Untereiner. 2004. Multiple mating results in multiple paternity in Richardson’s 
Ground Squirrels, Spermophilus richardsonii. Canadian Field-Naturalist 118(1): 90-94. 


Microsatellite DNA primers developed from Columbian Ground Squirrels (Spermophilus columbianus) were used to establish 
paternity in a Manitoba population of Richardson’s Ground Squirrels (Spermophilus richardsonii). Primers resolving 
variation at six microsatellite loci allowed ascription of paternity to 32 of 85 offspring born among litters of 15 breeding 
females sampled. While the failure to unambiguously document paternity for all juveniles precludes the use of these data to 
address questions of sperm competition and male mating success, the results do provide direct evidence that multiple mating 
by female Richardson’s Ground Squirrels results in multiple paternity within litters. 


Key Words: Richardson’s Ground Squirrel, Spermophilus richardsonii, mating, microsatellite DNA, multiple paternity, Manitoba. 


Differences in gamete size, and hence the differen- 
tial investment made in gametes by males and female 
ultimately define the sexes (Parker et al. 1972) and have 
promoted the evolution of disparate reproductive strate- 
gies. Males commonly enhance their fitness by obtain- 
ing copulations with more than one female (Bateman 
1948), maximizing the propagation of like copies of 
their genes by siring as many offspring as possible. 
Conversely, the enhancement of female fitness is typi- 
cally achieved through choosiness, whereby females 
mate only with males providing the greatest access to 
resources (Verner and Willson 1966; Thornhill 1976; 
Stanford 1995) and/or who are the most fit and con- 
tribute good genes to the female’s progeny (Zahavi 
1975; Weatherhead and Robertson 1979; Hamilton 
and Zuk 1982). At a proximate level, anisogamy also 
promotes sex-differential mating behaviour. The rela- 
tively few, large, nutrient-rich eggs produced by females 
necessitate only a single male ejaculate containing 
vast numbers of small, nutrient-poor spermatozoans to 
achieve fertilization, and thus female reproductive suc- 
cess may not increase through multiple mating. 

Despite these fundamental predictions following 
from anisogamy, multiple mating by females is com- 
mon in animals and can prove beneficial to females for 
various reasons. Obtaining multiple male mates may 
ensure fertilization (Hoogland 1998), enhance access 
to resources (Gray 1997), promote parental care on the 
part of males (Davies et al. 1996), confuse males as to 
paternity and thus reduce the probability of infanti- 
cide (Hrdy 1977), increase female fitness by ensuring 
genetic compatibility between the female’s eggs and 
at least some of the sperm (Zeh and Zeh 1996) or in- 
cite sperm competition (Gomendio and Roldan 1993), 
whereby the fitness of the female’s offspring is en- 
hanced by virtue of the good genes contributed by the 
victorious spermatozoans (Evans and Magurran 2000). 


Where multiple mating occurs, it is critical to un- 
derstand its implications to both males and females. 
Advances in molecular biology have revolutionized 
the study of reproductive behaviour, and revealed that 
observations of male/female association and mating 
behaviour may not be representative of parentage in 
free-living animal populations (Quinn et al. 1987). 
Methods for determining parentage (see Fleischer 1996 
for a review) have also become more powerful and ac- 
cessible, thereby providing researchers with an oppor- 
tunity to obtain comparative data which address both 
the proximate and ultimate questions pertaining to re- 
productive behaviour. 

Such a comparative picture has begun to emerge for 
the ground-dwelling squirrels (Spermophilus spp.), 
where multiple mating commonly results in multiple 
paternity within litters, and the first male to mate with 
a given female often sires a disproportionate number of 
offspring within a litter. Hanken and Sherman (1981) 
reported that 78% of 27 Belding’s Ground Squirrel 
(S. beldingi) litters for which paternity could be deter- 
mined using polymorphic blood proteins were multiply 
sired by between two and three males. Similarly, 50% 
of eight Thirteen-lined Ground Squirrel (S. tridecem- 
lineatus) litters were sired multiply, with the first male 
to mate siring 75% of offspring in those litters (Foltz 
and Schwagmeyer 1989). Boellstorff et al. (1994) de- 
tected multiple paternity in 88.9% of nine litters of 
California Ground Squirrels (S. beecheyi) and Murie 
(1995) reported multiple paternity in 15.8% of 165 
Columbian Ground Squirrel (S. columbianus) litters. 
Murie (1995), however, predicted that his data likely 
underestimated the incidence of multiple paternity 
given that only five polymorphic loci were used, and 
only one of those had more than two alleles. Indeed, 
using a more comprehensive and variable series of 
microsatellite primers (Stevens et al. 1997), multiple 


90 


2004 


paternity was detected in 64% of litters born to fe- 
male Columbian Ground Squirrels that had mated mul- 
tiply (Stevens, Strobeck and Murie unpublished data). 
Of those litters, 77% were sired by two males, 21% 
by three males, and 2% by four males. Further, there 
was a distinct first-male advantage, with 65% of the 
offspring sampled from among 77 litters sired by the 
first male to mate, and a progressively declining pro- 
portion of paternity success for subsequently mating 
males (Stevens, Strobeck, and Murie unpublished data). 

Despite the widespread occurrence of multiple pater- 
nity and a first-male mating advantage, deviations from 
that pattern are apparent among ground squirrels. 
Using both protein isozymes and DNA fingerprinting 
(see Burke 1989), Lacey et al. (1997) revealed that 
multiple paternity among Arctic Ground Squirrels (S. 
parryi) was relatively rare; it occurred in only | (9%) 
of 11 litters examined and the first male to mate sired 
the majority (90%) of the offspring in that litter. A 
departure from the first-male mating advantage has 
been detected in Idaho Ground Squirrels (S. brun- 
neus), Where Sherman (1989) reported a last-male 
mating advantage (a minimum of 66 — 100% of pups 
sired by the last/longest attending male) in the five of 
seven litters (71.4%) where multiple paternity was evi- 
dent. Additional data for other Spermophilus species 
will prove useful in determining whether these species 
are unique in their apparent departures from the gen- 
eral pattern, and thus how these expressions of multiple 
mating and sperm precedence impact the behaviour 
and fitness of males and females. 

Richardson’s Ground Squirrels (S$. richardsonii) 
are locally abundant throughout grazed areas of the 
Great Plains (Michener and Koeppl 1985). Females 
mate multiply (Michener and McLean 1996), and al- 
though this may result in multiple paternity within lit- 
ters (van Staaden et al. 1994, Michener personal com- 
munication), direct evidence of multiple paternity is 
lacking. Such data would contribute to our understand- 
ing of reproductive behaviour, and open new avenues 
for research into the contributions of paternal kinship 
to social interactions (e.g., Holmes and Sherman 1982) 
and patterns of dispersion among relatives in nature. 
We applied primers developed by Stevens et al. (1997) 
to prospective parents and offspring from litters of 15 
breeding females in a Manitoba population of Rich- 
ardson’s Ground Squirrels to test whether multiple 
mating results in multiple paternity within litters. 


Methods 
Study site, research subjects, and sample collection 
Research was conducted between 8 April and 26 
July 2002 on a 1.5-ha section of a larger 5-ha cattle 
pasture near Westbourne, Manitoba (50°10.190'N, 
98°38.103'W). Richardson’s Ground Squirrels on that 
site were live-trapped in National or Tomahawk traps 
baited with peanut butter and were marked for per- 
manent identification with numbered metal ear tags 


HARE, TODD, AND UNTEREINER: MULTIPLE MATING Q| 


(Monel #1 fish fingerling tags). Each squirrel was 
also given a distinctive mark on its dorsal pelage with 
black hair dye (Clairol Hydrience 52 Black Pear]). 
Trapping was conducted during the mating season (12 
through 25 April 2002) to determine mating dates of 
females and to monitor the reproductive condition of 
all squirrels. After mating had ended, trapping con- 
tinued to assess gestational progress in females, and 
a 2-mm tissue punch was taken from one pinna of each 
adult (10 males, 18 females). Samples were placed in 
1.5-ml microcentrifuge tubes in 95% ethanol, and 
stored at 4 — 7°C prior to DNA extraction and subse- 
quent molecular analysis. Fifteen females for which 
mating and parturition dates were known, and for whom 
ear punches had been obtained, and nest-burrow en- 
trances located were chosen to provide litters for pater- 
nity analysis. All juveniles in those litters were trapped 
within three days of their first emergence from their 
natal burrow, marked with ear tags and dye marks, and 
tissue sampled as described for adults. Animal research 
was conducted in accordance with the guidelines for the 
use of animals in research set forth by the Canadian 
Council on Animal Care. 


Molecular analysis of paternity 

Individual ear punches were placed in 1.5-mL micro- 
centrifuge tubes, centrifuged in a speed-vac for 5 min 
to remove the excess ethanol and digested at 55°C 
for 4 — 6 hrs in 70 uL of a sterile cell lysis solution 
(10 mM Tris-HCL, 100 mM EDTA, 2% SDS, pH 
8.0) containing 3 — 4 uL Proteinase K (10 mg/mL). 
Following the addition of 4 uL of RNase A, the tubes 
were mixed by inversion and incubated at 37°C for 
30 min. Proteins were precipitated by adding 23.5 uL 
of 7.5 M ammonium acetate to each tube of cell lysate 
and placing the samples on ice for 30 min. Tubes were 
centrifuged at 14000 rpm for 3 min and the superna- 
tant was transferred to a new 1.5-mL tube containing 
70 wL isopropanol. Samples were mixed by inversion, 
stored at -20°C for several hours and centrifuged at 
14000 rpm for 2 min. Following the removal of the 
supernatant, the pelleted DNA was washed in 75 UL 
of ice-cold 75% ethanol and centrifuged at 14000 rpm 
for 1 min. The ethanol was removed with a pipette 
and the tubes containing the DNA were placed in 
speed-vac for 5 — 10 min. The dried pellets were re- 
hydrated in 40 uL of sterile TLA buffer (10 mM Tris- 
HCL, 0.1 mM EDTA, pH 8.0) and stored at -20°C. 

Amplifications were performed on a GeneAmp® 
PCR System 9700 (Applied Biosystems, Foster City, 
CA) in a 25-uL volume containing ~ 200 ng DNA, 
0.8 mM dNTP, PCR buffer (0.1 M Tris-HCI pH 8.3, 
0.5 M KCl, 15 mM MgCl, 0.005 g gelatin/10 mL 
buffer), 0.5 units of Taq polymerase, and 0.2 UM con- 
centrations of each of seven primers (GS3, GS12, GS14, 
GS20, GS22, GS25, GS26) described by Stevens et al. 
(1997). The cycling conditions were 94°C for 2 min 
followed by two cycles of 94°C for 30 s, 58°C for 20 s, 
72°C for 5 s, and 33 cycles of 94°C for 15 s, 54°C for 


92 THE CANADIAN FIELD-NATURALIST 


20 s and 72°C for 5 s followed by a final extension at 
72°C for 30 s. PCR products were diluted (1:1) with 
sterile water and | uL of each diluted product was 
resolved by polyacrylamide gel electrophoresis on an 
ABI 373 Automated Sequencer (Applied Biosys- 
tems). Microsatellites were visualized and sized using 
GeneScan® 3.1 and Genotyper® 2.1 software (Applied 
Biosystems). 


Results 

Of the seven primers employed, one (GS22) result- 
ed in multiple banding patterns and was excluded 
from subsequent analysis. The remaining six primers 
resolved meaningful variation among individuals and 
were used in establishing familial relationships among 
the individuals sampled. Based upon the observed fre- 
quencies of the three to six alleles evident at those six 
loci (Table 1), the probability that two randomly drawn 
unrelated individuals will be identical at all loci is | 
in 9577 (from equation in Patekau and Strobeck 1994), 
Further, the probability that an unrelated male will be 
excluded as sire given knowledge of the mother’s geno- 
type is 0.91 (from equation in Chakravarti and Li 1983). 
All calculations assume that the loci examined are 
not linked, and provide highly conservative estimates 
of exclusion given that they are based on pooled data 
from dams, sires, and their offspring. 

Paternity was definitively ascribed to at least one 
juvenile within litters of all 15 dams included in the 
study. In total, however, unique identification of the 
male sire was achieved for only 32 of the 85 juveniles 
that emerged in those litters. Based upon those iden- 
tifications, multiple sires were detected in 12 of the 
15 litters examined (80%), with a minimum of two 
sires in 11 and a minimum of three sires in one of 
those 12 litters (2.1 + 0.1 sires/litter; mean + SE). 


Discussion 

The microsatellite primers developed by Stevens et 
al. (1997) proved effective in resolving multiple pater- 
nity within litters of Richardson’s Ground Squirrels. 
Indeed, with multiple sires in 80% of the 15 litters 


Vol. 118 


examined, multiple paternity is at least as common in 
Richardson’s Ground Squirrels as it is in other Spermo- 
philus species with the exception of California Ground 
Squirrels. 

Definitive genetic evidence of multiple paternity in 
the present study suggests that the use of the primers 
described by Stevens et al. (1997) in future studies of 
the reproductive and social behaviour of Richardson’s 
Ground Squirrels would prove quite fruitful. In concert 
with data on mating behaviour of the sort presented 
in Michener and McLean (1996), such studies could 
address the outcome of sperm competition and its rela- 
tion to mating order (Sherman 1989), the role of aggres- 
sion and male territoriality in affecting male mating 
success (Schulte-Hostedde and Millar 2002), and the 
factors underlying mate choice such as the effects of 
symmetry (Penton-Voak et al. 2001), body size 
(Gwynne 1981), paternal effort (Huber et al. 2002), 
and genetic relatedness (Bateson 1983; Chapman and 
Crespi 1998). The use of these microsatellite loci to 
assess relatedness among individuals (Queller et al. 
1993) would also allow insight into the role paternal 
kinship plays in influencing social behaviour (Widdig 
et al. 2002). Additional markers, however, would be 
necessary to comprehensively document the paternity 
of each and every individual. Microsatellite primers 
that were not employed in the present study, such as 
GS17 and GS34 (Stevens et al. 1997) or the 13 Idaho 
Ground Squirrel microsatellite primers described by 
May et al. (1997), would likely prove useful in this 
regard. 

The alleles identified in our Manitoba population 
(Table 1) differ from those described for the Richard- 
son’s Ground Squirrels examined by Stevens et al. 
(1997) in evaluating the efficacy of their primers on 
other Sciurids (see Table 2 in Stevens et al. 1997). 
Given that geographic variation, the primers employed 
here should also prove useful for studies of dispersal 
(Chapuisat et al. 1997), interpopulation differences 
related to conservation efforts (Patekau and Strobeck 
1994), and perhaps even in refining the taxonomy of 
the ground-dwelling squirrels (Gill and Yensen 1992). 


TABLE 1: Characteristics of microsatellite loci (from Stevens et al. 1997) applied in paternity analysis of Richardson’s Ground 
Squirrels. Results are based upon allele frequencies from 10 adult male, 15 adult female, and 85 juvenile squirrels. Expected 
heterozygosity and probability of identity were calculated according to formulae presented in Patekau and Strobeck (1994), 
and probability of exclusion was calculated following the methods of Chakravarti and Li (1983). 


Locus Alleles 

GS3 2215229523 18233 

GS12 147, 149, 152, 154 

GS14 242, 244, 246 

GS20 220; 222.237 

GS25 138, 142, 144, 146, 148, 154 
GS26 109, 111, 113, 115 


Expected Probability Probability 

heterozygosity of identity of exclusion 
0.630 0.178 0.347 
0.672 0.128 0.416 
0.549 0.298 0.253 
0.105 0.803 0.053 
0.769 0.057 0.557 
0.480 0.336 0.232 


2004. HARE, TODD, AND UNTEREINER: MULTIPLE MATING 93 
Acknowledgments Fleischer, R. C. 1996. Application of molecular methods to 
We are grateful to Curtis Strobeck and Lindsey the assessment of genetic mating systems in vertebrates. 
Carmichael of the University of Alberta for making Pages 133-161 in Molecular Zoology: Advances, Strategies, 
Fa and Protocols. Edited by J.D. Ferraris and S. R. Palumbi. 


the primers available to us and providing guidance 
on the use of the Genotyper® software, respectively. 
Gail Michener facilitated our work by allowing Mag- 
dalena Kujath to obtain samples from her study pop- 
ulation that proved useful in piloting the application 
of this technique. Our understanding of microsatellite 
methodology was enhanced through conversations 
with Nate Lovejoy, Ross McGowan, and Jan Murie, 
while Andrea Angus provided sound advice and assis- 
tance in the lab. We thank Jennifer Cassin, Kristin 
Melnyk, Melanie Sharpe, and David Wilson for capable 
field assistance and the Johnson family of Westbourne, 
Manitoba, for allowing us access to the squirrels on 
their land. Funding for this work was provided in part 
by a stipend from Human Resources and Development 
Canada in the form of a Summer Career Placement 
grant for Kristin Melnyk, and by the Natural Sciences 
and Engineering Research Council of Canada in the 
form of a University Summer Research Award to 
Glenda Todd, and operating grants to James Hare and 
Wendy Untereiner. Research infrastructure was also 
provided by a Canada Foundation for Innovation Res- 
earch Development Fund award to Wendy Untereiner. 
This is paper number 301 of the Delta Marsh Field 
Station. 


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Received 29 May 2003 
Accepted 15 June 2004 


Demographic Patterns and Limitation of Grey Wolves, Canis lupus, 
in and Near Pukaskwa National Park, Ontario 


S. ANNE FORSHNER!?, PAUL C. PAQUET?, FRANK G. M. Burrows*, GRAHAM K. NEALE, KEITH D. WADE®. 
and WILLIAM M. SAMUEL 


'Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9 Canada 
*Mailing address: P.O. Box 10, Jasper National Park, Jasper, Alberta TOE 1E0 Canada 

Faculty of Environmental Design, University of Calgary, Calgary, Alberta T2N 1N4 Canada 
4Bruce Peninsula National Park, Tobermory, Ontario NOH 2RO Canada 

‘Garcia and Associates, Bozeman, Montana 59718 USA 

°Pukaskwa National Park, Heron Bay, Ontario POT 1RO Canada 


Forshner, S. Anne, Paul C. Paquet, Frank G. M. Burrows, Graham K. Neale, Keith D. Wade, and William M. Samuel. 2003. 
Demographic patterns and limitation of Grey Wolves, Canis lupus, in and near Pukaskwa National Park, Ontario. 
Canadian Field Naturalist 118(1): 95-104. 


In response to concern regarding the growth and long-term viability of the wolf population in and near Pukaskwa National Park, 
a study of demographic patterns and limitation of radio-collared wolves (Canis lupus) was completed between 1994 and 1998. 
The mean annual finite rate of increase (0.96) suggested that population growth of wolves was limited and declining slightly. 
Small pack sizes, high cumulative mortality, and low reproductive success also suggested a declining population. Two limiting 
factors, ungulate biomass and human-caused mortality, were examined to determine the importance of each in limiting the 
population growth of wolves. Ungulate biomass was involved because occurrence of natural-caused mortality was high (9 of 17 
wolves) compared with other studies. In addition, consumption rates were low and similar to other studies where starvation 
and other signs of malnutrition were noted. Further, Moose densities in the study area were low to moderate and below thresholds 
indicating nutritional stress for wolves. Occurrence of human-caused mortality was high (8 of 17 wolves) suggesting that it was 
also an important limiting factor, particularly given the low availability of ungulate biomass and reproduction noted in this study. 


Based on present demographic patterns, ungulate biomass, and human-caused mortality, the wolf population likely will remain 


at present low densities or continue to decline. 


Key Words: Canis lupus, wolves, limitation, demographic patterns, Pukaskwa National Park, Ontario. 


In 1996, researchers in the Greater Pukaskwa Eco- 

system (GPE) (Figure 1) in Ontario postulated that two 
limiting factors were negatively affecting the growth 
and long-term viability of the Grey Wolf (Canis lupus) 
population (Burrows et al. 1996"). These limiting fac- 
tors were low availability of ungulate prey and high 
mortality from human causes. Managers in Parks Can- 
ada Agency were concerned because wolves are a 
native species in the GPE. Thus, Parks is legislated to 
protect and further, to ensure the long-term viability of 
that population (Parks Canada 2000). This prompted 
a study of wolves in the western half of the GPE, in- 
_ cluding Pukaskwa National Park (PNP). 
In this paper, we quantify population limitation of 
_ wolves and examine the importance of ungulate bio- 
mass and human-caused mortality in limiting the popu- 
lation growth of wolves, 1994-1998. For these pur- 
_ poses, we review and discuss demographic data on wolf 
_ densities, population growth, reproduction, and mortal- 
ity. Further, we report and discuss data on the avail- 
ability of prey and rates of kill and consumption. 


' * References marked with asterix (*) are listed in a separate 
Documents Cited section following Acknowledgements, 
all others are in Literature Cited. 


Study Area 

The study area comprised 4500 km? in the western 
half of the GPE on the north shore of Lake Superior in 
Ontario (48°N and 85°W) (Figure 1). The area includes 
PNP (1878 km?) but also adjacent land with intensive 
forestry, gold mines, towns and associated infrastruc- 
ture. 

Two distinct physiographic regions, coastal and inte- 
rior, occur within the study area. The coastal region is 
characterized by rugged topography with elevations 
ranging from 189 to 650 m. Many lakes and rivers occur 
in the area, creating a patchy landscape. The interior 
region is a flat plateau characterized by a heavily erod- 
ed landscape of mountains previously scoured by con- 
tinental glaciers (Poitevin et al. 1989"). 

Mean annual precipitation is 74 cm along the coast 
and 64 cm inland. Winter and summer temperatures 
range from —13C° — 14.6C° for the coastal area and 
—17C° — 15.9C° inland (Poitevin et al. 1989"). Ice cov- 
er on Lake Superior ranges annually from 5-100% 
(Skibicki 1994’). 

Vegetation on the coast along Lake Superior and in- 
land is mixed with associations of Balsam Fir (Abies 
balsamea), Jack Pine (Pinus banksiana), White Birch 
(Betula papyrifera), White Spruce (Picea glauca), 
Black Spruce (Picea mariana), Eastern White Cedar 


95 


96 THE CANADIAN FIELD-NATURALIST 


Lake 
Superior 


Legend 


| __ Greater Pukaskwa 
Ecosystem 
M8 Pukaskwa National Park 
wolf study area 
Highway 17 N 
Peete earier en eaten 
: 20 km M 
|Prepared by: Parks Canada 


a | Pukaskwa National Park 


2001 —_L. Parent 


Vol. 118 


FicureE |. Location of the wolf (Canis lupus) study area within the Greater Pukaskwa Ecosystem, Ontario, Canada (center 


48°N, 85°W). 


(Thuja occidentalis), and Trembling Aspen (Populus 
tremuloides), with occasional Red Maple (Acer ru- 
brum) and other hardwoods more locally abundant in 
the southeastern corner of the study area. 

Predatory mammals included Grey Wolf, Black Bear 
(Ursus americanus), Red Fox (Vulpes vulpes), Lynx 
(Lynx canadensis), River Otter (Lontra canadensis), 
Fisher (Martes pennanti), American Marten (Martes 
americana), Mink (Mustela vison), and Weasels (Mus- 
tela spp.). Coyotes (Canis latrans) were rare except 
around towns. 

Moose (Alces alces) were the primary and most 
abundant ungulate prey species for wolves. Woodland 


Caribou (Rangifer tarandus tarandus) were few and 
concentrated in small bands along the coast of Lake 
Superior (Bergerud 1985). Numbers ranged from 6-14 
in PNP, 1993-1997 (Wade 1993*, 1995*, 1997", 1999*). 
White-tailed Deer (Odocoileus virginianus) were rare 
in the GPE. 


Methods 
Capture and handling 

We attempted to locate wolves in as many packs as 
possible. Wolves were captured with modified leg- 
hold traps in summer (7 = 21) and by using a net-gun 
from a helicopter in early winter (n = 5). All wolves 


| 


2004 


Legend 
A Heron Bay 


@ Mobert First Nation 

© Marathon 

tx White River 
see22 Swallow River Pack 
—-—- Bremner River Pack 
—--— White River Pack 
== Cascade Lake Pack 
—-— Neys Pack 


|_| Rein Lake Pack 
— — Black River Pack 
(Cee Be} 


0 _20 km 


| Prepared by: Parks Canada 
te Pukaskwa National Park 
2001 _L. Parent 


iiaebn 
* 
° ~ 


2 


annneeeed 


FORSHNER ET AL.: GREY WOLVES IN PUKASKWA NATIONAL PARK 97 


i Highway 17 


Ficure 2. Annual (1 April — 31 March) home ranges of wolves in the study area, 1994-1998. Home ranges are 95% MCP. 
Years for home ranges of packs were Swallow and Bremner River Packs, 1997-1998; White River Pack, 1996-1998; 
Cascade Lake Pack, 1995-1996; Neys Pack, 1995-1997; Rein Lake Pack, 1994-1998; Black River Pack, 1994-1996, 


1997-1998. 


were immobilized with Telazol® (tiletamine hydro- 
chloride (HCL) and zolazepam HCL, A.H. Robins 
Co., Richmond, Virginia). Rectal temperature, pulse, 
and respiration of wolves were closely monitored 
throughout the procedure. Immobilized wolves were 
examined for injuries, equipped with conventional 
VHF transmitters (Lotek®, Newmarket, ON), weighed, 
sexed, and aged. A committee for care of wild animals 
approved all capture and handling operations (Wildlife 
Animal Care Committee, Ontario Ministry of Natur- 
al Resources, 1994-1997). 


Biotelemetry 

The target frequency for locating each radio-collared 
wolf was four times/month in summer (April-Octo- 
ber) and six to eight times/month in winter (Novem- 
ber-March). Wolves were located by plane using a 
portable receiver (Lotek® SRX-400), right-left switch 
boxes, and paired three-element Yagi antennae mount- 
ed on the wing struts of a fixed-wing aircraft (Cessna 
185). Wolf location was recorded with a Global Posi- 
tioning System (Garmin® 75 Aviation). For all loca- 
tions, transmitter frequency, observer, date, time of 
location, number of wolves, color of wolves, and pre- 


sence of pups were recorded. Mean error of telemetry 
(difference between observed and true location) was 
calculated by using data we collected when regularly 
locating stationary transmitters placed throughout the 
study area. Location data were downloaded into a Geo- 
graphical Information System (GIS [Tydac SPANS*]) 
for display and analysis of wolf movements. 


Home Ranges 

Ranges V® software (Kenward and Hodder 1996°) 
was used to calculate annual (1 April — 31 March) 
sizes of home ranges. To represent these areas we 
used relocations of packs and 95% minimum convex 
polygons (MCP) (Mohr 1947). All obvious extrater- 
ritorial forays and dispersals were excluded from the 
analyses (Ballard et al. 1997). We assumed home 
ranges were defined when the observation-area curve 
formed an asymptote (Kenward and Hodder 1996") 
and locations were obtained throughout the year. 

For each pack we used one radio-collared wolf/year 
to represent the annual home range of the pack. This 
is reasonable as locations from one wolf indicate 
position of the entire pack when a high degree of 
association exists among pack members (Kolenosky 


98 THE CANADIAN FIELD-NATURALIST 


and Johnston 1967; Fuller and Keith 1980; Fritts and 
Mech 1981; Ciucci et al. 1997). This condition was 
confirmed in this study by aerial observations of 
packs during telemetry flights. 

Accuracy of locations for the entire study was 150 m, 
which was the mean error of telemetry obtained by 
all participants. Accordingly, we changed the fix reso- 
lution from the Ranges V® default of 1 m to 150 m. 
This resolution was used to set the width of the bound- 
ary strip that was included in polygon edges and 
areas (Kenward and Hodder 1996"). We left the scal- 
ing parameter at the software default of 1 m, which 
means that each coordinate unit was | m from the next. 


Density, pack sizes, and population growth 

Density of wolves/1000 km? was calculated by 
determining intra-pack densities (number of wolves 
in pack/home range size) and averaging these den- 
sities/year (Potvin 1987; Bjorge and Gunson 1989; 
Okarma et al. 1998). The number of wolves in a pack 
was based on the maximum number of wolves ob- 
served in mid-winter (15 January-15 February). We 
defined a pack as a group of two or more wolves that 
traveled together for more than one month (Messier 
1984). In two cases we had insufficient data to deter- 
mine the sizes of home ranges, thus we followed Mes- 
sier (1985) and used data from previous or subsequent 
years. 

Population growth or the mean annual finite rate of 
increase was calculated based on the ratio of succes- 
sive yearly estimates of density (Fuller 1989). 


Reproduction 

We did not observe wolves at dens during this study. 
Dense vegetation and the secretive nature of wolves 
precluded accurate visuals of wolf groups until Octo- 
ber or November, at which time pups were difficult to 
distinguish physically from adults. Hence, successful 
year-specific reproduction was ascertained when: (1) 
pups were captured in spring; or (2) a pack increased 
in size from March to the following December, pro- 
viding that sites of focal activities (e.g., pup-resting 
areas) were observed in the intervening time (Messier 
1985). Unsuccessful reproduction (i.e., no or failed 
reproduction) was ascertained when: (1) a pack did 
not demonstrate focal activity sites in the summer; or 
(2) a pair remained together from March to the follow- 
ing December (Messier 1985). Results are reported for 
each pack by year. 


Moose density 

To examine availability of ungulate biomass to 
wolves, we used Moose density (moose/1000km?) 
based on aerial surveys using stratified random sampl- 
ing (Gasaway et al. 1986*). More specifically, from 
1993 to 1999 a single Moose density was calculated for 
PNP and the three Wildlife Management Units (21A, 
21B, 33) surrounding PNP where wolf packs were 
distributed. There was little or no change in Moose 


Vol. 118 


density among yearly estimates (Burrows 2001), thus 
we averaged results from 1993-1999 for each area. 


Rates of kill and consumption of prey by wolves 

The rates of killing and consumption of large prey 
by wolves in four packs were studied by daily aerial 
and ground observation, January-March 1998. The 
Bremner River Pack was located 57 times between 
18 January and 27 March 1998 (69 days) and the 
Rein Lake Pack was located 57 times between 8 Janu- 
ary and 26 March 1998 (79 days). Other packs locat- 
ed were the White River Pack, 22 times between 11 
February and 20 March 1998 (38 days) and the Swal- 
low River Pack, 22 times between 9 February and 22 
March 1998 (42 days). To calculate the kill rate, we 
recorded the number of animals killed by wolves/ 
tracking period and the number of wolves present at 
the kill (Messier 1985). Prey killed were located 
from the air and from ground-based tracking. At kill 
sites, we confirmed prey species, time, and cause of 
death. For only the largest pack of wolves (Bremner 
River), in addition to aerial locations, we simultane- 
ously snow-tracked wolf movements and collected 
scats to determine if all kill sites were found with the 
aerial telemetry. Technicians at Big Sky Laboratory 
(PO Box 0776, Florence, Montana 59833-0776) iden- 
tified prey remains by macroscopic examination and 
comparison with known material and hair-scale im- 
pressions (Adorjan and Kolenosky 1969°). 

In this analysis we considered only tracking sessions 
where pack locations were not separated by >54 hr. 
There were a few exceptions, however, where loca- 
tions were separated by 72 hr. These periods were 
retained in the analysis because wolves made a kill or 
visited one of several garbage dumps the day they 
were relocated making it unlikely that we missed a 
kill. Nonetheless, kill rates in this study should be 
considered minimums as wolves were not relocated 
every day and some small prey such as deer (fawns 
and adults), Caribou calves, Beaver, and other smaller 
prey items may have been missed. It is unlikely, how- 
ever, that we missed many of these kills because 
White-tailed Deer and Caribou were rare in the study 
area. We report kill rates as ungulates killed/wolf/day. 

Rates of consumption were calculated based on 
kill rates and average weights of wolves and prey. We 
calculated the whole weight of wolves based on the 
average from radio-collared adults and other wolves 
found dead in the study area. The average edible 
weights of Moose and beaver prey were assumed to 
be 330, 261, 114, and 13 kg for adult male Moose, 
adult female Moose, young-of-the-year Moose and 
Beaver, respectively (Peterson 1977; Thurber and 
Peterson 1993). We assumed the average weight of a 
White-tailed Deer was 105 kg for an adult male (Kolen- 
osky 1972; Forbes and Theberge 1996). Eighty % of 
the adult deer carcass was considered edible (Pimlott 
1967; Forbes and Theberge 1996). All consumption 
rates are reported as kg prey/kg wolf/day. 


2004. FORSHNER ET AL.: GREY WOLVES IN PUKASKWA NATIONAL PARK 99 
TABLE 1. Sizes of annual home ranges and home range areas/wolf of seven packs of Grey Wolves (Canis lupus) in the study 
area, 1994-1998. 
Early Late 
winter winter 
number number 100% 95% Number 
of wolves of wolves MCP MCP of radio Area/wolf 

Year Pack in pack* in pack? (km?)° (km?)° fixes (km?)¢ 
1994-1995 Black River® 6 4 283 156 53 26 

Rein Lake 3 3 310 249 38 83 
1995-1996 Black River® 4 4 548 388 5 97 

Rein Lake 3 3 600 533 58 178 

Neys* 3} 4 269 244 96 8] 

Cascade Lake! 1 204 170 26 170 
1996-1997 Rein Lake® 2 | 561 Syi/ 39 279 

Neys* 6 4 113 101 96 17 

White River 2 2 407 345 Sy) 173 
1997-1998 Black River® 4 3 468 450 74 113 

Rein Lake® 2 1 692 600 87 300 

White River 2, 2 589 498 65 249 

Bremner River 6 3 760 644 93 107 

Swallow River 5 3 567 500 62 100 

Mean 8}55) DEG 455 388 63 139 

SE 0.5 0.3 52 48 6.3 25 


“Maximum pack size, 15 January-15 February. 
> Maximum pack size, March. 


© Sizes of home ranges were described using the minimum convex polygon method (MCP) (Mohr 1947). 


‘Based on 95% MCP. 
© Packs that used town dumps. 


‘Home range size and home range area/wolf were not included in mean because the areas were not fully defined; i-e., the 
observation-area curve was asymptotic but locations were not obtained throughout the year. 


Mortality and survival of radio-collared wolves 

We completed survival analysis for radio-collared 
wolves from 20 August 1994 to 31 December 1998. 
Wolves were re-located from time of capture until 
mortality or the radio-signal disappeared. For known 
deaths we estimated the date of mortality to the near- 
est day using evidence from the field. When evidence 
was unavailable, day of mortality was deemed the 
midpoint of the interval between the last day the wolf 
was known alive and the day it was discovered dead. 
The cause of mortality was often identified on site 
and when possible, confirmed by necropsy. 

We calculated the cumulative mortality of radio- 
collared wolves (n = 25) using the Kaplan-Meier prod- 
uct limit estimator and Minitab (Version 12) software. 
One of 26 captured wolves was shot by a trapper 
while in the research trap and is not included in the 
analysis. Cause of mortality was described using %. 
We assumed the proximate cause of death was the 
ultimate cause of death. We were unable to assess the 
relative importance of other factors that may have 
been involved. 


Results 

Twenty-six adult wolves were captured and then 
radio-collared (n = 25) or tagged (n = 1) from 1994- 
1997. These animals represented seven packs and 
one lone wolf. Two of seven packs occurred almost 
exclusively in the park and all wolf packs were radio- 
collared in the study area. There were no other wolf 
packs in the study area during this study. We followed 
two packs in 1994-1995, four in 1995-1996, four in 
1996-1997, and six in 1997-1998. The average mass 
of adult female wolves (n = 11) was 26.9 + 1.4 kg 
and that of adult males (n = 14) was 36.5 + 2.8 kg. 


Home ranges 

Sizes of annual home ranges (Figure 2) of seven 
packs across 13 pack-years were adequately described 
in this study (Table 1). Estimates accurately repre- 
sented areas used by wolves because sizes of annual 
home ranges were not correlated with number of 
relocations (r, = 0.52, 0.05 > P > 0.02). Home range 
sizes of packs and home range areas/wolf were vari- 
able. The average annual home range size was 388 + 
SE 48 km2 (95% MCP, n = 13, range = 101-644 km’) 


100 


(Table 1). The average home range area/wolf was 
139 + SE 25 km? (95% MCP, n = 13, range = 17-300 
km?) (Table 1). 


Density, pack sizes, and population growth 

Wolf density did not change over time; recorded 
densities were 7.9, 9.6, and 7.2 wolves/1000 km? in 
1995-1996 (n = 4 packs), 1996-1997 (n = 4), and 1997- 
1998 (n = 6), respectively. Density declines, however, 
if the Neys pack (Figure 2) is excluded from the 
calculations. This pack exclusively used dumps for 
food (Krizan 1997) and the home range was much 
smaller compared with all other packs in the study 
(Table 1). Accordingly, wolf densities were 7.1, 5.9, 
and 5.9 wolves/1000 km? in 1995-1996 (n = 3 packs), 
1996-1997 (n = 3), and 1997-1998 (n = 5). 

Average mid-winter (15 January — 15 February) 
pack size was 3.5 + SE 0.5 wolves (n = 14 pack-years) 
(Table 1). This average declined in late winter (March) 
to 2.7 + SE 0.3 (n = 14 pack-years). The number of 
wolves in all except two packs remained stable or 
declined, 1994-1998. Numbers fluctuated annually in 
the Neys and Swallow River Packs (Table 2). Accord- 
ingly, the mean annual finite rate of increase from 
1995-1998 was 0.96. 


Reproduction 

From spring 1994 to spring 1998, wolves reproduced 
successfully in eight of 22 pack-years (36%) (Table 3). 
This was a maximum estimate of successful repro- 
duction. In two of eight pack-years, we assumed wolves 
had reproduced because large numbers of wolves were 
noted in the packs in the following early fall and winter. 


Moose density 

Average densities of Moose varied among manage- 
ment units. Management Unit 33 had the highest 
Moose density (0.285 + 0.03-0.07 moose/km? 90% CI) 
followed by Unit 21A (0.225 + 0.02-0.03), 21B 
(0.220 + 0.02-0.03), and PNP (0.153 + 0.03-0.08). 


Rates of kill and consumption 

The Swallow River (3 wolves) and Bremner River 
Packs (5) killed and consumed more ungulates than 
the White River (2) and Rein Lake Packs (2). The kill 
rates for each pack respectively were 6.8, 3.4, 0.0, and 
0.0 ungulates/wolf/day. Consumption rates were 0.21, 
0.11, 0.0, and 0.0 kg prey/kg wolf/day. The White 
River and Rein Lake Packs did not kill any ungu- 
lates; however, both packs scavenged from various 
sources. The White River Pack scavenged from Moose 
that were killed by vehicles or trains, from other wolf 
kills, and from snare sets. The Rein Lake Pack scav- 
enged from refuse in the town dump for White River 
(Figure 2). 


Mortality and survival of radio-collared wolves 

As of 31 December 1998, 17 of 26 wolves radio- 
collared or tagged from 1994 to 1998 were dead, only 
four were confirmed alive, and five were missing. Eight 
wolves died from human causes: trains killed three, 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE 2. Highest numbers of wolves in packs in the study 
area, 1994-1998. The number of sightings is in parentheses. 


Pack size* 
Pack 1994 1995 1996 1997 1998 
Rein Lake 3(4) = 3(3) 1(2) 1> 0 
White River D (2) ee (A) 2(2) 2(9) 2(2) 
Bremner River 3(3)P 
Cascade River 1(4) 0 
Black River 4Q) VAC) w4e@) 3(4) 
Swallow River 5 (D)o 3(3) 4(1) 
Neys 3(4) — 4(2) 2(2)) 4) 


* Maximum numbers of wolves seen in March. 
> Pack sizes were confirmed by track-counts made from 
the ground. 


TABLE 3. Reproductive success of wolf packs, 1994-1998. 


Year 

Pack 1994. 1995 1996 1997 =1998 
Black River +4 4 - + + 
White River “pb = = = = 
Rein Lake +b = = = = 
Neys + - ? 
Bremner River ~ + 
Swallow River = = 
“+ = reproduced successfully; — = did not reproduce 


successfully. 
> This pack likely reproduced successfully because many 
wolves (10) were seen in late fall (November — December). 


three were snared, and two shot. Nine wolves died 
from natural causes: two starved, two were killed by 
other wolves, four died from disease (three from mange 
and one from blastomycosis), and one died from un- 
known natural causes. We assumed this last wolf was 
not killed by humans because we were in a remote 
area, there were no signs of humans in the area, and 
we found no bullets, snares, or other human devices. 
Survival of radio-collared wolves decreased between 
one and three years post-collaring. Wolves had a 32% 
(SE 0.10) chance of dying in the first year, a 30% (SE 
0.15) chance of dying in the second year, and a 57% 
(SE 0.26) chance of dying in the third year. Median 
survival time was 689 days or 1.9 years post-collaring. 


Discussion 
Population limitation of wolves 

The growth rate of the wolf population in the study 
area was limited from 1995-1998. The mean annual 
finite rate of increase, 0.96, indicated a 4% rate of 
decline. This rate of increase is not unique, however, 
and similar rates recorded from other populations 
have varied from 0.93-2.40 (Theberge and Strickland 
1978; Fritts and Mech 1981; Ballard et al. 1987; 
Hayes et al. 1991"; Messier 1991; Pletscher et al. 1997). 


\ 


| 


i 
{ 


2004 


In addition to the estimated rate of growth, there are 
a number of other factors that suggest the wolf popula- 
tion was declining slightly. First, pack sizes were small 
and generally declining. Mean pack size (3.5 wolves + 
SE 0.5) was much smaller than the average of 10 wolves 
for packs that hunt moose in North America (Mech 
1970). Furthermore, five of seven packs in this study 
remained stable or declined in size from 1994 to 1998. 
If this population were increasing in size, the number 
of wolves within packs would likely increase. This 
happened in the Yukon where rapid increases in pack 
sizes of colonizing wolves were the primary means by 
which an intensively reduced wolf population reached 
their pre-reduction densities (Hayes and Harestad 
2000). 

High cumulative mortality of wolves is the second 
factor that suggested a declining population. We com- 
pared the cumulative rate of mortality from the first 
year (32%) of our study with annual rates of mortal- 
ity from other studies. There is little agreement among 
researchers on the annual rate of mortality that causes 
a population decline in wolves. However, Fuller (1989) 
reviewed several wolf studies across North America 
and concluded that populations would stabilize with 
an overall annual mortality rate of 35%. Given this, it 
appears the mortality rate in this study was sufficient 
to account for the slightly declining rate of growth in 
this study. 

Coupled with high mortality of adult wolves, low 
reproductive success of wolves in this study suggested 
a population decline. Wolves reproduced successfully 
in only 36% of possible occasions compared with 
45-93% noted in other areas (Messier 1985; Potvin 
1987; Peterson et al. 1998; Hayes and Harestad 2000). 
We were unable to determine if wolves produced pups 
that died soon after birth or whether whelping occurred 
at all. Lack of denning, however, suggests no pups 
were produced. 


Limiting factors 

We examined the importance of two factors that lim- 
ited the growth of the wolf population in this study: un- 
gulate biomass and human-caused mortality. Ungulate 
biomass is commonly reported to limit growth of other 
wolf populations (Mech 1977a; Fuller and Keith 
1980; Packard and Mech 1980; Keith 1983; Messier 
1985; Peterson and Page 1988) and data from this 
study suggest it was important. The strongest data rep- 
resented the occurrence of natural-caused mortality. 
In this study, more than half (9 of 17) of radio- 
collared wolves died from natural causes, which is high 
compared with other North American studies (Peter- 
son 1977; Carbyn 1982; Peterson et al. 1984; Ballard 
et al. 1987; Hayes et al. 1991”). Starvation and intra- 
specific aggression were responsible for four of nine 
(24%) wolves dead in this study and have been report- 
ed common in other populations where ungulate bio- 
mass is low. For instance, in southwestern Quebec, 


FORSHNER ET AL.: GREY WOLVES IN PUKASKWA NATIONAL PARK 


10] 


Messier (1985) noted wolves with fewer prey available 
incurred more deaths from natural causes, namely 
starvation and intraspecific aggression. In that area, 


similar to our study area, moose density was 0.23 
moose/km? and there were no other ungulate species 
present. Similarly, Mech (1977a) noted occurrence of 
starvation and intraspecific aggression increased as 
prey availability declined in Minnesota. 

Disease was the other natural cause of death ob- 
served in this study. Four of nine wolves (24%) died 
from either sarcoptic mange or blastomycosis. Blasto- 
mycosis is enzootic in Minnesota (Schlosser 1980) and 
Wisconsin (Sarosi et al. 1979; McDonough and Kuzma 
1980) but until now (Krizan 2000; Paquet et al. 2001), 
had not been reported from other wolf populations 
across North America. This level of disease-related 
mortality has not been reported in any other popula- 
tions of wolves. In other populations, disease accounts 
for 2-21% of wolf mortality (Carbyn 1982; Peterson 
et al. 1984; Fuller 1989; Ballard et al. 1997) and is 
often not even reported (Messier 1985; Ballard et al. 
1987; Potvin 1987; Hayes et al. 1991°; Meier et al. 
1995; Pletscher et al. 1997). The only other study 
where disease clearly affected a wolf population was in 
Alaska where rabies accounted for 21% of wolf mor- 
tality and was a significant factor in the decline of the 
population (Ballard et al. 1997). Disease cannot be 
linked with certainty to low ungulate biomass but wolves 
that lack food should be more vulnerable to disease than 
those with more food available. Furthermore, food 
shortage leading to nutritional stress could combine 
with disease factors to increase the significance of 
otherwise innocuous or sub-lethal conditions (Brand 
et al. 1995). 

We also examined rates of consumption of ungu- 
late prey to determine the importance of ungulate bio- 
mass as a limiting factor. Consumption rates for three 
of four packs in this study were low (Bremner River, 
Rein Lake, and White River). These packs consumed 
<0.13 kg/kg wolf/day, which Mech (1977b) deter- 
mined is the minimum rate of consumption required 
for all wolves to survive and rear pups successfully. 
Two packs killed no ungulates and relied on scav- 
enging to survive (White River and Rein Lake Packs). 

These data suggest that at least three of four packs 
could have been limited by food. Indeed, in the White 
River Pack, the dominant female failed to reproduce 
the following spring and was extremely emaciated 
(mass = 23.5 kg) when killed by other wolves later in 
the summer. Similarly, the Rein Lake Pack was reduced 
to one wolf by winter 1997. She did not reproduce 
the following summer and to survive, she scavenged 
mainly from the town dump for White River (Figure 
2). She was dead as a result of mange by December 
1998 (mass = 28.5 kg). The Bremner River Pack may 
have been limited by food but data were not strong. 
One wolf dispersed in summer 1998 and died from 
unknown natural causes. Other wolves could have dis- 


102 


persed and died later because pack numbers dropped 
from nine to three over the winter 1997-1998 (un- 
published data). Some members of this pack, however, 
did survive and reproduce two years in a row. 

Most of the rates of consumption in this study are 
similar to those from other areas where starvation and 
other signs of malnutrition of wolves were noted. For 
instance, Messier (1987) noted more deaths of wolves 
from malnutrition in areas of low density of moose 
(0.23 moose/km7) where wolves had 0.05 kg/kg wolf/ 
day (based on kill rate of 1.7 kg/wolf/day and wolf 
mass of 32.3 kg). Peterson and Page (1988) noted 
starvation and other indicators of severe nutritional 
stress in an area of high Moose density (1.9 moose/ 
km?) when food availability fell below 0.12 kg/kg 
wolf/day (based on kill rate of 4.0 kg/wolf/day and 
wolf mass of 32.3 kg). 

As a final method to assess the importance of un- 
gulate biomass, we examined density of Moose, the 
main prey for wolves in this study. Moose density 
was low to moderate (0.153-0.285 moose/km?) and 
similar to Moose densities in other areas where wolves 
were nutritionally stressed. Messier (1987) found that 
in areas where Moose density dropped below 0.4 
moose/km*, wolves were nutritionally stressed. He 
also reported that below 0.2 moose/km? wolf packs 
could not subsist and (or) reproduce successfully 
(Messier 1985). 

The second limiting factor we examined was human- 
caused mortality. Besides ungulate biomass, it is the 
other most commonly reported factor that limits the 
growth of wolf populations (Gasaway et al. 1983; 
Keith 1983; Peterson et al. 1984; Fuller 1989; Noss 
et al. 1996; Paquet et al. 1996"). In other areas where 
human-caused mortality was considered the primary 
limiting factor, it accounted for 69-80% of all mortal- 
ity (Peterson et al. 1984; Ballard et al. 1987, 1997). 
In our study area, human causes accounted for only 
47% of mortality of adult radio-collared wolves. None- 
theless, human-caused mortality is likely still impor- 
tant, particularly given the low ungulate biomass and 
reproduction noted in this study. Gasaway et al. (1983) 
found that in areas with low ungulate biomass, har- 
vest levels as low as 20% can limit wolf populations. 
Fuller (1989) found that wolf populations with low 
productivity can withstand less overall mortality be- 
cause there are fewer pups, which often make up dis- 
proportionate amounts of harvests. 

In conclusion, the population growth of wolves in 
this study area was limited and declined slightly based 
on (i) mean annual finite rate of increase; (ii) small and 
generally declining pack sizes; (iii) high cumulative 
mortality; and (iv) low reproductive success. 

Based on these demographic patterns, low availabil- 
ity of ungulate biomass and existing levels of human- 
caused mortality, this population likely will remain at 
present low densities or continue to decline. This situa- 
tion is challenging to managers for Parks Canada Agen- 
cy because the study area, which includes a National 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Park, may not have a highly productive source popu- 
lation for wolves. Further, protection for wolves out- 
side the park is limited because few restrictions exist 
regarding the nature, timing, and extent of wolf har- 
vesting. 


Acknowledgments 

Field research was funded by Parks Canada. We 
thank all PNP wardens and staff for field and techni- 
cal support, particularly K. Wade, S. Sutton, and C. 
Strong. Additionally L. Parent provided much GIS 
expertise. Thanks also to J. Whittington and R. Whit- 
tington for field support and pilots W. Roberts and 
M. Robb for many safe hours of flying time. Thanks 
to Helicopter Wildlife Management Team for skillful 
capturing and collaring of animals. All animal treat- 
ment procedures were approved by the Wildlife 
Animal Care Committee, Ontario Ministry of Natural 
Resources (Permit Numbers 1994-13, 1995-13, 1996- 
13, 1997-13). We thank J. Whittington, B. Dobson, 
and M. Boyce who incisively reviewed later drafts of 
this manuscript. S.A. Forshner was personally support- 
ed by an NSERC scholarship, the University of Alber- 
ta, a University of Alberta Teaching Assistantship, 
Bill Samuel, and Parks Canada. 


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Received 13 March 2003 
Accepted | March 2004 


Estimation of Seed Bank and Seed Viability of the Gulf of Saint 
Lawrence Aster, Symphyotrichum laurentianum, (Fernald) Nesom 


Joni F. Kemp and CureisTIAN R. LACROIX! 


Department of Biology, University of Prince Edward Island, 550 University Avenue, Charlottetown, Prince Edward Island 
C1A 4P3 Canada 
'Author to whom correspondence should be addressed. 


Kemp, Joni F., and Christian R. Lacroix. 2004. Estimation of seed viability of the Gulf of Saint Lawrence Aster, Symphyotrichum 
laurentianum (Fernald) Nesom. Canadian Field-Naturalist 118(1): 105-110. 


The Gulf of St. Lawrence Aster, Symphyotrichum laurentianum, is a member of the family Asteraceae and is listed as “threatened” 
by COSEWIC (Committee on the Status of Endangered Wildlife in Canada). This rare and vulnerable halophyte grows in only 
a few locations in New Brunswick, Prince Edward Island, and the Magdalen Islands, Quebec. As an annual, S. laurentianum 
relies exclusively on its seeds to survive to the next generation. The goal of this study was to estimate the quantity of viable 
S. laurentianum seeds in the persistent and transient seed banks at selected sites in Prince Edward Island. Overall, the number 
of seeds in the transient and persistent seed banks is low. The greatest concentration of seeds was found near the surface of 
the soil. In addition, only a small proportion of those seeds tested positive for viability based on Tetrazolium staining. Of the 
seeds in the persistent and transient seed banks combined, 53% were viable whereas only 2% of the seeds in the persistent seed 
bank were viable. Population surveys were also completed at the five known sites (both extinct and extant) in Prince Edward 
Island National Park. All sites showed signs of decline based on population estimates dating back to 1993. The Covehead 
Pond site showed the greatest decline: from 250-300 individuals in 1993 to only 10 individuals in 2002. The population at 
Dune Slack also showed a dramatic decrease from approximately 65 000 in 1999, to 2 200 individuals in 2002. Monitoring 
of this plant and the development of a management plan for the species are critical to its survival. 


Key Words: Symphyotrichum laurentianum, Aster laurentianus, Gulf of St. Lawrence Aster, seed viability, Tetrazolium, 


transient and persistent seed banks, rare plant, COSEWIC, Prince Edward Island National Park. 


The Gulf of St. Lawrence Aster, Symphyotrichum 
laurentianum (Fernald) Nesom, is a member of the 
family Asteraceae and is listed as “threatened” accord- 
ing to COSEWIC (Committee on the Status of Endan- 
gered Wildlife in Canada 2004*). It is an annual plant 
that relies exclusively on its seeds to survive until the 
next germination season, which makes this plant very 
vulnerable. S. /aurentianum is a rare halophyte only 
found in a few areas in Prince Edward Island (PEI), 
New Brunswick (NB), and the Magdalen Islands in 
Quebec. It grows in salt marshes that are only flooded 
during spring and high tides. The St. Lawrence Aster 
has smooth, hairless, and fleshy linear-lanceolate to 
spatulate, generally sessile leaves (Figure 1). It can grow 
up to 30 cm tall and can bear | to over 700 flower heads 
(Houle and Haber 1990). Each flower head is subtended 
by leafy bracts and is composed exclusively of white to 
pink disk florets surrounded by a white pappus (Stewart 
2000* and Figure 1). Seeds are enclosed in achene fruit 
walls when they are dispersed. 

In 1990 fewer than 1000 plants were known to exist 
in PEI, NB and Quebec (Magdalen Islands) combined. 
However, further research and surveys have produced 
several new sites. While several sites in PEI and Que- 
bec contain more than 1000 individuals each, the St. Law- 
rence Aster is still vulnerable to extinction (Gilbert et 
al. 1999*; Stewart and Lacroix 2001). Recent research 
conducted by Stewart (2000*) showed that population 
sizes could suddenly decline dramatically. 


The extent of the persistent seed bank of a plant must 
be known in order to fully understand the germina- 
tion ecology of the species (Baskin and Baskin 1998). 
In addition, the viability of a plant’s seeds must also 
be maintained in order for the plant to be successful 
(Fenner 2000). A seed bank is a reserve of viable, un- 
germinated seeds in the soil. Seeds are considered vi- 
able if they have the ability/potential to germinate under 
favourable conditions (Baskin and Baskin 1998). There 
are two categories of seed banks: transient and persis- 
tent. A transient seed bank refers to a short-term storage 
of seeds, that is seeds that are viable for one year or one 
germination season. The seeds in the transient seed bank 
therefore germinate in the first year after dispersal. Any 
seeds that are dormant and survive to subsequent ger- 
mination seasons are considered to be part of the per- 
sistent seed bank. The persistent seed bank contains 
seeds that remain viable in the soil for at least two ger- 
mination seasons, making this a long-term reserve of 
seeds that are at least one year (or one germination sea- 
son) old. A knowledge of both the extent of the seed 
banks and the germination potential of seeds of S. lau- 
rentianum constitutes crucial baseline data to develop 
a management plan for the species. 

The specific goals of this study were: (1) to estimate 
the quantity of seeds in the seed bank (both transient 
and persistent) for S. lJawrentianum at each of the select- 
ed sites in PEI in order to evaluate the ability of the 
plant to maintain current populations, and (2) to test the 


105 


106 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


FiGurE |. Representative photographs of Symphyotrichum laurentianum, pre-flowering (A) and post-flowering (B). 


viability of seeds in the persistent seed bank and the 
transient seed bank. 


Materials and Methods 
Population Surveys 
Five sites within the PEI National Park (Figure 2) 
. Were visited. Total population counts were made at the 
Covehead Pond and East Marsh A sites because of the 
small number of plants at these locations. The Western 
Wetland site was also surveyed to confirm that no 
asters were growing there. Population estimates were 
recorded for Dune Slack and East Marsh B sites be- 
cause of the large population sizes. These estimates 
were accomplished by using a grid system. The gener- 
al area where the plants were growing was subdivided 
into 0.5 m? sections. A random number of sections was 
selected and all plants within these grids were counted. 
These counts along with the total area of the site were 
used to extrapolate the approximate number of plants 
found at each site. 


Collection of Soil Samples 

Soil cores were taken from five sites within PEI 
National Park (Figure 2), the only location in the pro- 
vince where the plant is found: Covehead Pond, Dune 
Slack, East Marsh A, East Marsh B and Western Wet- 
land. Three transects were run at each site except Cove- 
head Pond. Twenty coring points were flagged along 
three transects in Dune Slack. Due to sampling limita- 
tions imposed by Parks Canada, only ten coring points 
were flagged along three transects at East Marsh A and 
a further ten along the three transects at East Marsh B. 
Similarly, ten coring points were also flagged along the 
three transects for Western Wetland. Eleven coring 
points were flagged at Covehead Pond in a slightly 


different manner because of the physical nature of the 
site. Four transects were run from the water’s edge of 
the pond to the high tide mark along the traditionally 
heavily populated side of the pond. Eight coring points 
were flagged evenly among these transects and the re- 
maining three points were spaced evenly along the other 
side of the pond where the plants were also reported 
to grow. 

Two sets of soil cores were taken from each site: one 
during late June/early July, and another during late 
August/early September. These dates were chosen to 
correspond to the type of seed bank that was sampled. 
The late June/early July samples were taken before the 
emergence of the 2002 season plants, to ensure that the 
transient as well as persistent seed banks were includ- 
ed. The late August/early September samples were 
taken after all the seeds that were going to germinate 
that season had germinated. Consequently, only the 
persistent seed bank was included in this specific sam- 
ple date. 

All the core samples were taken using an Oakfield 
“LS” 36” soil sampler with an effective coring length 
of 24 cm and an inner diameter of | inch. In order to 
study the vertical distribution of seeds in the soil, each 
soil core was divided equally into a top, middle, and 
bottom section. Each section of the core was labelled, 
placed into a separate bag, and put in a freezer set at 
-4°C until processing which took place two weeks to 
three months later. All seeds found in the soil samples 
were tested for viability. 

Viability Testing 
Seeds were tested for viability using 2,3,5-triphenyl 


tetrazolium chloride (TTC) (Grabe 1970). The TTC 
test is based on a colour reaction: any seeds that are 


2004 


KEMP AND LACROIX: SEED BANK AND SEED VIABILITY 107 


FIGURE 2. Map of all Prince Edward Island sites used in this study. Study sites are depicted as squares on both maps. All five sites 
shown in (A) are located in the national park: 1. Covehead Pond, 2. Western Wetland, 3. Dune Slack, 4. East Marsh B, 
and 5. East Marsh A. (B) Location of sites on Prince Edward Island with black square highlighting the specific area 


shown in (A). 


viable will stain pink/red. Complete achenes (contain- 
ing seeds) were placed on a piece of filter paper in a 
petrie dish, and left overnight to soak in distilled water. 
The next day, when the seeds were fully imbibed with 
water, they were dissected out of the achene fruit 
wall and seed coat. After removal of the seed coat, 
the embryos were placed in a 1% TTC solution for 
two hours at 35°C, as prescribed for dicotyledonous 
seeds. The embryos were then evaluated for viability 
according to the scheme developed in Grabe (1970). 
As a general rule, the radicle, shoot tip and cotyledons 
must stain pink/red in order for the seed to be con- 
sidered viable. 


Statistical Analysis 

The viability of seeds in the soil was compared 
graphically for vertical distribution and sampling dates. 
Statistical analysis was not possible on these data be- 
cause there was no replication. Due to the vulnerable 
status of the plant, sampling was limited by restrictions 
on collecting imposed by Parks Canada. The vertical 
distribution of seeds and fruit walls within the soil was 
compared using Kruskal-Wallis tests (non parametric 
equivalent of ANOVA). All data from the five sam- 
pling sites on PEI were pooled for analysis. 


108 


Results 
Population Sizes 

Population sizes for all known PEI sites are shown 
in Table 1. All populations experienced a decrease in 
numbers, the most noticeable of these being the pop- 
ulation at the Dune Slack site with more than 95% 
decline. The smallest population at the Covehead Pond 
site also experienced a decrease from 243 individuals 
in 1999 to only 10 individuals in 2002. 


Seed Banks 

A total of 122 cores were taken from PEI sites. Of 
these, 32 contained at least one seed. There were sig- 
nificantly more seeds in the top portion of those cores 
than in any other portion of the soil samples (Figure 
3; Kruskal-Wallis test; P < 0.05, n=45 for each por- 
tion). The non-random distribution of the seeds, as evi- 
denced by the highly variable number of seeds found 
in the core samples, is also highlighted in Figure 3. 
This may be related to where the plants clustered the 
year before, a factor we were not able to determine 
prior to establishing coring sites. 


Seed Viability 

There was a noticeable decline in the percentage of 
viable seeds among the top, middle, and bottom sec- 
tions of the soil cores taken on PEI. We found that 
28.4% of the seeds in the top section of the cores were 
viable compared to 20% of the seeds in the middle sec- 
tion of the cores. None of the seeds in the bottom section 
of the cores were viable. 

There was a marked decline in the percentage of via- 
ble seeds between the late June/early July sample and 
the late August/early September sample (Figure 4). 
The percentage of viable seeds in the persistent seed 
bank (2%) is practically non-existent when compared 
to the viability of the seeds in the persistent and tran- 
sient seed banks combined (53%). This means that the 
majority of viable seeds germinate within one year of 
being produced. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Discussion 
Population Sizes 

Symphyotrichum laurentianum populations on PEI 
continue to show dramatic fluctuations and it can be 
very difficult for an annual plant to rebound from these 
declines in population size. Although the Dune Slack 
site was considered one of the two largest populations 
on PET]; its rate of decline, if maintained, will bring the 
population down to a size comparable to the smallest 
populations. A comparatively small population at 
Covehead Pond has been relatively stable at 150 to 
300 individuals for the past ten years but the last popu- 
lation count found only ten individuals which, cou- 
pled with the low viability of seeds in the seed banks, 
indicates that this site is on the verge of extinction if 
preventive measures are not taken. 

The Island Nature Trust, a local conservation orga- 
nization, conducted a survey of suitable areas on PEI 
in 2002 to locate new populations of S. /aurentianum. 
Several prime candidate habitats were located but un- 
fortunately no asters were found at those sites (Mac- 
Quarrie, personal communication). 


Seed Banks 

There appears to be a lack of mixing of the soil in 
the salt marshes which causes a clustering of seeds in 
the top sections of the soil. This trend is obvious in the 
vertical distribution of seeds shown in Figure 3. Seeds 
that remain trapped in the upper layer of the soil are 
more likely to get exposed to the elements thereby 
potentially affecting the viability of the embryo. 

One of the most meaningful results from this study 
is the fact that the viability of seeds in the persistent 
seed bank is practically non-existent. This can be dev- 
astating for an annual plant such as S. laurentianum. 
Without the long-term storage of viable seeds, this plant 
is vulnerable to short term changes in its habitat, both 
natural and anthropogenic. 


TABLE |. Comparison of population sizes for Symphyotrichum laurentianum on Prince Edward Island. 


Population GPS Location 


Covehead Pond 46° 25' 49.39" N 


63° 09' 07.98" W 


46° 24' 47.88" N 
63° O1' 16.89" W 


Western Wetland 


Dune Slack 46° 24' 51.49" N 
62° 59' 41.98" W 
East Marsh A 46° 24' 55.97" N 
62° 58' 43.89" W 
East Marsh B 46° 24' 55.27" N 
62° 59' 06.11" W 


* estimate 


1992 1999 2002 
168 243 10 
425 0 0) 
15 000 — 20 000* 65 250* 2 200* 
133 
50 000 — 60 000* 25 000 — 60 000* 
(Sites A and (Sites A and 44 100* 


B combined) B combined) 


| 
) 


2004 


40 ; 
2 
Qa. 
5 
om 30 
th, 
o 
jo 
gat ° 
io ‘0 
o 
ep) 
— * 
2 mn 
o 
a 
E =— 

yea 
Top Middle Bottom 


Portion of the Core 
(N= 32) 


FiGuRE 3. Box plots showing median (centre line) and 25" and 
75" percentile for numbers of seeds found in cores 
from all Prince Edward Island soil samples that con- 
tained seeds. Outliers (*) are included to show the 
non-random distribution of the seeds. 


During the summer of 2002, the Dune Slack site was 
heavily flooded during a storm. This caused the site, and 
consequently the maturing aster plants, to be smoth- 
ered in eel grass. Since the Dune Slack population had 
already experienced a dramatic decrease in size between 
1999 and 2002, the necessary number of mature viable 
seeds needed to survive this natural short-term event 
may not be available. 

Covehead Pond is an example of a site that is very 
likely to become extinct due to anthropogenic causes. 
This site is isolated from Covehead Bay by a road. The 
culvert that is presently in place did not allow proper 
drainage from the pond during the summer of 2002 and 
caused this site to be flooded. The ten plants that were 


100 a 


n 

xo} 

o 

o 

no 

o 

a 60 

& 

> 

es 

fo} 

oO 

RB 40 4 

= 

oO 

2 

a) 

ae 20 | 

) 
i) 1 2 3 
Persistent and Transient Persistent 
N=89 N=93 


Seed Bank Type 


FiGurE 4. The percentage of viable seeds in the seed banks on 
Prince Edward Island. Persistent and transient seed 
banks = all late June/early July samples on Prince 
Edward Island. Persistent seed bank = all late August/ 
early September samples on Prince Edward Island. 


KEMP AND LACROIX: SEED BANK AND SEED VIABILITY 


109 


seen at this site in 2002 were flooded by water and also 
smothered by eel grass. This occurred at a crucial time 
during seed maturation. Only one plant was found at 


the Covehead site in 2003. 


Status of the Species 

In 2003, there were 441 Canadian species at risk, 
and 140 of these were plants (Committee on the Status 
of Endangered Wildlife in Canada 2004%*). Until 
recently, there has been very little in terms of Federal 
legislation in place to protect these species. The Species 
at Risk Act (SARA) which came into effect in 2002 will 
hopefully facilitate this process especially with regard 
to developing a plan for the recovery of every extir- 
pated, endangered, or threatened species. 

The results of this study show a disturbing trend: the 
estimated number of seeds in the seed banks is low, the 
viability of the seeds in those seed banks is also low 
(especially in the persistent seed bank), and the popu- 
lations are declining. The Gulf of St. Lawrence Aster 
is currently under review and it is important that all 
available information on the biology of the species be 
considered (especially information pertaining to seed 
banks). The status of this plant has recently been up- 
dated to “threatened” which is defined as “a species 
that is likely to become endangered if nothing is done 
to reverse the factors leading to its extirpation or ex- 
tinction” (Committee on the Status of Endangered 
Wildlife in Canada 2004*). This new designation con- 
firms the urgent need to design a management plan for 
S. laurentianum to ensure that it does not become ex- 
tinct. Although harsh weather patterns leading to strong 
winds and flooding cannot be prevented (i.e., extreme 
flooding of the Dune Slack site during summer of 
2002), alternate measures such as transplanting ex situ 
plants into affected sites should be considered. Pre- 
liminary results show that seedlings of S. laurentianum 
can be grown from seed and transplanted from cul- 
ture medium to soil with a relatively high success 
rate (Stewart 2000*). However, a population decline 
and loss of suitable habitat due to anthropogenic 
activities can be prevented and further steps. such as 
ensuring proper drainage at a site such as Covehead 
Pond, should be taken. 


Acknowledgments 

This research was partially funded by an NSERC 
(National Sciences and Engineering Research Council) 
operating grant to C. Lacroix. We would like to thank 
Parks Canada for allowing us access to the sites to 
collect samples. Thank you to Kate MacQuarrie for 
her comments on this manuscript. 


Documents Cited (marked * in text) 

Committee on the Status of Endangered Wildlife in Can- 
ada. 2004. COSEWIC assessment and update status report 
on the Gulf of St. Lawrence Aster, Symphyotrichum lauren- 
tianus in Canada. Ottawa. 39 pages. Retrieved May 2004, 
froma http://www.cosewic.gc.ca 


110 


Gilbert, H., J. Labrecque, and J. Gagnon. 1999. La situa- 
tion de l’aster du Saint-Laurent (Aster laurentianus, syn. : 
Symphyotrichum laurentianum) au Canada. Ministére de 
l'Environnement Direction de la conservation et du patri- 
moine écologique, Québec. 

Stewart, S. E. 2000. Micropropagation of Aster lauren- 
tianus. Unpublished honours thesis, University of Prince 
Edward Island, Charlottetown, Canada. 


Literature Cited 

Baskin, C. C., and J. M. Baskin. 1998. Seeds: Ecology, bio- 
geography, and evolution of dormancy and germination. 
Academic Press, San Diego. 

Fenner, M. 2000. Seeds: The ecology of regeneration in 
plant communities. CABI Publishing, New York. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Grabe, D. E., Editor. 1970. Tetrazolium testing handbook for 
agricultural seeds: contribution Number 29 to the hand- 
book on seed testing. Tetrazolium Testing Committee of 
the Association of Official Seed Analysts. 

Houle, F., and E. Haber. 1990. Status of the Gulf of St. Law- 
rence Aster, Aster laurentianus (Asteraceae), in Canada. 
Canadian Field-Naturalist 104: 455-459. 

Stewart, S. E., and C. R. Lacroix. 2001. Germination poten- 
tial, updated population surveys and floral, seed and seed- 
ling morphology of Symphyotrichum laurentianum, the 
Gulf of St. Lawrence Aster, in the Prince Edward Island 
National Park. Canadian Field-Naturalist 115: 287-295. 


Received 2 June 2003 
Accepted 3 June 2004 


Consumption of Shrews, Sorex spp., by Arctic Grayling, 


Thymallus arcticus 


JONATHAN W. Moore! and G. J. KENAGY? 


' Department of Biology, University of Washington, Box 351800, Seattle, Washington 98195 USA; 


e-mail: jwmoore @u. washington.edu 


* Burke Museum and Department of Biology, University of Washington, Box 353010, Seattle, Washington 98195 USA 


e-mail: kenagy @u.washington.edu 


Moore, Jonathan W., and G. J. Kenagy. 2004. Consumption of shrews, Sorex spp., by Arctic Grayling, Thymallus arcticus. 


Canadian Field-Naturalist 118(1): 111-114. 


In an investigation of the dietary habits of Arctic Grayling (Thymallus arcticus) we found that two individuals out of 93 sampled 
in southwestern Alaska (approximately 59°N, 159°W) contained a total of five shrews (Sorex spp.). These shrews contained 
enriched levels of nitrogen stable isotopes, suggesting utilization of nutrients derived from salmon. We hypothesize that 
normally terrestrial shrews accidentally enter streams while foraging along the productive riparian zones of creeks with high 
densities of salmon. Shrews are apparently susceptible to opportunistic predation by resident stream fishes, including Arctic 


Grayling, when they enter the streams. 


Key Words: Arctic Grayling, Thymallus arcticus, diet, salmon-derived nutrients, shrews, Sorex spp., stable isotopes. 


The Arctic Grayling (Thymallus arcticus) is a com- 
mon nearctic freshwater fish in lakes and streams at 
high latitudes (Nelson 1994). Grayling typically feed 
on a variety of aquatic and terrestrial invertebrates in 
the water column or air-water interface (Armstrong 
1986). In addition, Arctic Grayling occasionally supple- 
ment their diets with salmon eggs and small fish 
(Armstrong 1986). Despite existing knowledge of Arctic 
Grayling biology and its regional importance, the ecol- 
ogy and behavior of this species remain not well un- 
derstood, including the extent to which Arctic Grayling 
exploit terrestrial sources of prey (Armstrong 1986). 

As part of an extensive study of the aquatic and asso- 
ciated terrestrial ecosystem and its relationship to sal- 
mon in southwestern Alaska (Schindler et al. 2003), 
we had the opportunity to investigate dietary habits of 
Arctic Grayling. We discovered evidence of apparently 
rare consumption by Arctic Grayling of small terres- 
trial mammals of the Order Insectivora, shrews of the 
genus Sorex. 

During summer 2001 we sampled resident fishes in 
five streams that flow into Lake Nerka in the Wood 
_ River-Tichick State Park, Dillingham County, south- 
western Alaska (approximately 59°36’N, 159°05’W), 
by conventional and fly angling. The study streams sup- 
| port large runs of Sockeye Salmon (Oncorhynchus 
_ nerka). Captured fish were anesthetized, marked with 
subcutaneous tags, weighed, measured, and released. 
_ We obtained diet samples by back-flushing the stomach 
with a gastric lavage. Diet samples were preserved in 
| ethanol and transported back to the University of 
Washington for analysis. 

_ We sampled the stomach contents of 93 Arctic Gray- 
| ling from five tributaries of Lake Nerka. Two of the 
93 Arctic 2001 Grayling contained visible evidence of 
the body parts of shrews (Figure 1, Table 1). One Arctic 


Grayling (length 394 mm, mass 591 g), captured on 
13 August in the Little Togiak River contained the 
remains of three shrews. A second Arctic Grayling 
(length 391 mm, mass 522 g), captured on 25 August 
in Elva Creek contained the remains of two shrews. 
Nothing else about the two observed Arctic Grayling 
was unusual other than their stomach contents. Although 
the fish were relatively large, they were not extreme, 
being longer than about 70% of the other Arctic Gray- 
ling. In addition to the shrews, the diets of these two 
fish contained aquatic and terrestrial insects typical of 
the diets of the other Arctic Grayling sampled at this 
same time and location. 

The five alcohol-preserved shrews (Sorex spp.) from 
the diet samples of two Arctic Grayling were deposit- 
ed in the Burke Museum mammal collection. Three 
shrews (UWBM 74151, 74152, 74154) were sufficient- 
ly intact that we could examine individual teeth in the 
skull, which is critical for species identification. We 
used the keys of Nagorsen (1996), based on body and 
skull measurements and dental characteristics, particu- 
larly the relative length of the third unicuspid. In two 
cases, digestion had progressed too far to allow species 
determination by these characters. We subsequently 
referred tissue samples of all five specimens to Eric 
Waltari, Idaho State University, for molecular (cyto- 
chrome-b) identification, according to protocols used 
in his systematic investigation of Sorex, as developed 
by Demboski and Cook (2001). Molecular identification 
confirmed the three initial morphological identifica- 
tions and provided definitive identification for the two 
previously unidentifiable individuals (UWBM 74153, 
74155) 

The two shrews from Elva Creek were identified as 
Sorex monticolus and Sorex cinereus (Figure 1, upper). 
The S. cinereus specimen had a wet weight of 3.3 g 


IU 


112 


and was fairly intact except for digestion of the fur 
away from the skin. Due to more extensive digestion, 
the S. monticolus specimen could not be measured and 
weighed accurately, but its intact skull and dentition 
allowed a positive identification (Table 1). 

The three shrews from the Little Togiak River (Fig- 
ure 1, lower) included a highly intact S. cinereus with 
considerable fur still attached to its skin (Figure 1, 
lower, center). The other two shrews were extensively 
digested and could only be identified by molecular 
analysis, as S. cinereus (left) and S. monticolus (right). 
In summary, the two Arctic Graylings from two dif- 
ferent streams both had fed on shrews of two species 
(Table 1). 

The presence of the shrews Sorex monticolus and 
Sorex cinereus in this region of southwest Alaska ly- 
ing to the north of Bristol Bay and east of the Kilbuck 
Mountains is consistent with the known geographic 
ranges of shrews (Hall 1981) and a more recent dis- 
tributional analysis (S. O. MacDonald and J. A. Cook, 
personal communication). Shrew abundance and diver- 
sity are high in this area, and three other species are 
also present: S. hoyi, S. tundrensis, and S. yukonicus. 

Although some species of the genus Sorex live and 
feed primarily in water (Beneski and Stinson 1987), 
none of the five Sorex species in the study area is pri- 
marily aquatic. However, all five occur to some extent 
in riparian habitat, where their success is enhanced by 
the aquatic component of the food chain. As a result 
of this habitat association, the shrews are potentially 
accessible to predatory fishes in these waters. The ten- 
dency for shrews of the genus Sorex to enter water, 
whether for foraging or dispersal, has been reviewed 
by Hanski (1986). In addition to intentional entry into 
the water, it is also possible that these shrews occa- 
sionally enter the water accidentally, resulting from 
their normal activities near the water’s edge. 

Shrews may be attracted to the riparian zones of 
these creeks because of the potential foraging oppor- 
tunities offered by abundant salmon carcasses. Ceder- 
holm et al. (1999) list shrews, including Sorex cinereus, 
as known scavengers of salmon carcasses. Evidence 
from stable isotopes ('°N) from our study (Table 1) 
and Ben-David et al. (1998) show that shrews (Sorex 
spp.) living near streams containing anadromous sal- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


mon runs bear the enriched isotopic signature of feed- 
ing directly or indirectly on salmon, for example, on 
insects that feed on salmon carcasses. We analyzed 
the stable isotope signature of muscle tissue from each 
shrew. Because anadromous salmon have a relatively 
high isotopic signature (6 °N = 11-13) compared to 
most other potential food sources, isotopes have been 
used to trace salmon nutrients through both aquatic 
and terrestrial food webs (Kline et al. 1990; Ben- 
David et al. 1998). The abundance of '°N in shrew 
muscle tissue was measured by combusting several 
mg of ground muscle tissue from each shrew in a mass 
spectrometer by the Stable Isotope Lab at the Univer- 
sity of California, Davis. Stable isotopes were expressed 
as a delta (6) value—the deviation from an isotope 
standard (atmospheric nitrogen). In other words, 
SION) = (GSNae Nie ee) ON reenact) XoLOOO! 

We used a two-sample t-test to compare 6°N of 
shrews in diets of Arctic Grayling to published values 
of isotopes from six shrews that were collected more 
than 500 m from streams with salmon in southeast- 
ern Alaska by Ben-David et al. (1998). The five 
shrews we collected from Arctic Grayling diets were 
relatively enriched, averaging 5.88 + 0.47 6 PN 
(mean + standard error; Table 1), higher than shrews 
collected from sites over 500 m from Alaskan 
salmon-bearing streams, that averaged 4.5 + 0.3 6 
!SN (ty9 = 2.24, P < 0.10) (Ben-David et al. 1998). 
This relative enrichment suggests that the shrews we 
found in the Arctic Grayling diets may have been 
feeding directly on salmon carcasses or indirectly, 
for example, by feeding on insects that had been 
feeding on salmon carcasses. The high productivity 
of these riparian zones, especially along streams with 
high densities of salmon, probably contributes posi- 
tively to the success and survival of shrews, the tiniest 
of mammals, in this extreme northern environment. 

The skewed distribution of the number of shrews 
contained per Arctic Grayling is evidence that con- 
sumption of shrews is not a random event. We assessed 
whether Arctic Grayling consumption of shrews was 
random by comparing the observed frequency distri- 
bution of the number of shrews in Arctic Grayling 
stomachs to the expected distribution, assuming pre- 
dation to be random. We calculated expected frequen- 


sample 


TABLE |. Characteristics of shrews removed from the stomachs of two Arctic Grayling. 


Sorex spp 6 ©N! Location 

S. cinereus? 4.73 Elva Creek 

S. monticolus* SP) Elva Creek 

S. monticolus* 6.33 Little Togiak River 
S. cinereus® 7.42 Little Togiak River 
S. cinereus* 5.70 Little Togiak River 


Body dimensions? Specimen number 


96-38-11 mm UWBM 74151 
indeterminable UWBM 74152 
indeterminable UWBM 74153 
88-38-12 mm UWBM 74154 
indeterminable UWBM 74155 


' 8I5N is the stable isotope of nitrogen and can be used to trace salmon-derived nutrients. 
? Dimensions are total length, tail length, and hind foot length, in mm. 


> Identified both by dentition and cytochrome-b analyses. 


“Identified by only cytochrome-b analysis because of missing/digested dentition. 


2004 


MoOoRE AND KENAGY: CONSUMPTION OF SHREWS BY GRAYLING 


FiGurRE 1. Shrews removed from the stomachs of two Arctic Grayling, in various stages of digestion. Upper: Shrews from a 
522 g Arctic Grayling sampled at Elva Creek (59°34’N, 159°05’W) — Sorex monticolus (UWBM 74152) left and 
Sorex cinereus (UWBM 74151) right. Lower: Shrews from a 591 g Arctic Grayling at Little Togiak River (59°36’N, 
159°04’W) — Sorex cinereus (UWBM 74155) left, S. cinereus (UWBM 74154) center, and Sorex monticolus 


(UWBM 74153) right. 


cy distributions using the Poisson function in Matlab 
5.313, assuming an average number of shrews per 
Arctic Grayling of 0.054 (5 shrews in 93 Arctic Gray- 
ling). We encountered fish with either two or three 
shrews in their stomachs, but no fish with only one 
shrew. Based on a random expectation for Arctic Gray- 
ling consumption, there was a 0.12 probability of sam- 
pling an Arctic Grayling with two shrews and only a 
0.003 probability for three shrews. Thus we conclude 
that Arctic Grayling consumption is not random, and 
we speculate that this is due to individual differences 
in either the effectiveness or motivation of individual 
Arctic Grayling as shrew predators or scavengers. It 
may also be due to differences in the probability of 
shrews entering the feeding sites of specific Arctic 
Grayling. 

Although Arctic Grayling are typically considered 
to be specialists on aquatic and terrestrial insects (Arm- 
strong 1986), our observation indicates that some 
Arctic Grayling are opportunistic feeders, capable of 
consuming small mammals. Shrews, typically 4-8 g 
(Nagorsen 1996), represent a meal for Arctic Graylings 
that provides a large amount of energy compared to 
that of typical individual invertebrate food items. Con- 
sumption by Arctic Grayling of small mammals, in- 
cluding shrews, has been reported by several previous 
authors. Alt (1978*) reported an unspecified small 
number of shrews in diets of Arctic Grayling in the 
Fox River of western Alaska. De Bruyn and McCart 
(1974*) found that seven out of 136 Arctic Grayling 
contained a single shrew in the Firth River, Yukon, 
Canada. A European study of the grayling Thymallus 


thymallus indicated a 10% incidence of small mam- 
mals in the diet, particularly the shrew Sorex araneus, 
and the capture of these fish at night by bait angling 
suggested their ability as nocturnal hunters of semi- 
aquatic prey such as shrews (Teplov 1943). Miller 
(1946) reported that one out of 102 Arctic Grayling 
at Great Bear Lake, Canada, contained juvenile lem- 
mings (rodent) of an unspecified species in the diet. 
Reed (1964*) also found juvenile lemmings in two 
Arctic Grayling out of 1300 individuals sampled in 
the Tanana River drainage, Alaska. 

Terrestrial food sources often subsidize the diets of 
freshwater fish, and these food sources can include 
small mammals (for example, Brown Trout [Salmo 
trutta], Cochran and Cochran 1999; and Largemouth 
Bass [Micropterus salmoides|, Hodgson and Kinsella 
1995). Although Arctic Grayling are traditionally con- 
sidered insectivores, our report demonstrates that they 
also opportunistically consume shrews that venture 
into freshwaters. 


Acknowledgments 

We thank the Alaska Salmon Project of University 
of Washington for logistic support and the National 
Science Foundation and ARCS foundation for finan- 
cial support. D. E. Schindler and M. D. Scheuerell 
provided insightful discussions and helped collect 
fish diets. M. D. Scheuerell and J. I. Jones prepared 
the shrew muscle samples fer isotopic analysis, which 
were run by D. Harris. We thank E. Waltari for molec- 
ular identification of the shrews and R. Rausch for 
advice on the observation and literature. 


114 


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de Bruyn, M., and P. J. McCart. 1974. Life history of the 
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Northwest Territories. Edited by P. J. McCart. Canadian 
Arctic Study, Ltd., Calgary, Biological Report Series 15. 
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Reed, R. J. 1964. Life history and migration patterns of Arc- 
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River drainage of Alaska. Alaska Department of Fish and 
Game Research Report Number 2: 8-30. 


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Beneski, J. T., and D. W. Stinson. 1987. Sorex palustris. 
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Cederholm, C. J., M. D. Kunze, T. Murota, and A. Sibatani. 
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Cochran, P. A., and J. A. Cochran. 1999. Predation on a 
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mouse, Mus musculus, by brown trout, Salmo trutta. 
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THE CANADIAN FIELD-NATURALIST 


Vol. 118 


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Hall, E. R. 1981. The mammals of North America. John 
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signifier Richardson, from Great Bear Lake. Copeia 1946: 
227-236. 

Nagorsen, D. W. 1996. Opossums, shrews, and moles of 
British Columbia. University of British Columbia Press, 
Vancouver. 169 pages. 

Nelson, J. S. 1994. Fishes of the World. Third edition. John 
Wiley and Sons, New York. 600 pages. 

Schindler, D. E., M. D. Scheuerell, J. W. Moore, S. M. 
Gende, T. B. Francis, and W. J. Palen. 2003. Pacific 
salmon and the ecology of coastal ecosystems. Frontiers 
in Ecology and the Environment |: 31-37. 

Teplov, V. P. 1943. The significance of the common shrew 
(Sorex araneus L.) and some other vertebrates in the nutri- 
tion of Thymallus thymallus L. [in Russian]. Zoological 
Journal 22: 366-368. 


Received 9 June 2003 
Accepted | June 2004 


Unusual Behavior by Bison, Bison bison, Toward Elk, Cervus elaphus, 
and Wolves, Canis lupus 


L. Davip Mecu!, Rick T. McINTYRE2? and DouGLAS W. SMITH? 


'U.S. Geological Survey, Northern Prairie Wildlife Research Center, 8711 - 37" Street, SE, Jamestown, North Dakota 58401 - 
7317 USA; Mailing address: The Raptor Center, 1920 Fitch Avenue, University of Minnesota, St. Paul, Minnesota 
55108 USA 

* National Park Service, Yellowstone Center for Resources, P. O. Box 168, Yellowstone National Park, Wyoming 82190 USA 

3 Mailing address: General Delivery, Silver Gate, Montana 59081 USA 


Mech, L. David, Rick T. McIntyre, and Douglas W. Smith. 2004. Unusual behavior by Bison, Bison bison, toward Elk, Cervus 
elaphus, and Wolves, Canis lupus. Canadian Field-Naturalist 118(1): 115-118. 


Incidents are described of Bison (Bison bison) in Yellowstone National Park mauling and possibly killing a young Elk (Cervus 
elaphus) calf, chasing wolves (Canis lupus) off Elk they had just killed or were killing, and keeping the wolves away for 
extended periods. During one of the latter cases, the Bison knocked a wolf-wounded Elk down. Bison were also seen approach- 
ing wolves that were resting and sleeping, rousting them, following them to new resting places and repeating this behavior. 
These behaviors might represent some type of generalized hyper-defensiveness that functions as an anti-predator strategy. 


Key Words: Bison, Bison bison, Wolt, Canis lupus, Elk, Cervus elaphus, Yellowstone National Park. 


Bison (Bison bison) are known to be aggressive to directed the capturing and radio-tagging of the Wolves, 
other ungulates including Elk and even to kill Elk allowing them to be located and observed. 
(Cervus elaphus) calves (Chapman 1937: 148; Rush 
1942: 225; McHugh 1958: 143; Mahan 1977). Little Observations 
is known about the causes and circumstances of this 1. 30 May 1995 — Bison Attack Elk Calf 
aggressiveness except for the scattered reports docu- A recently born Elk calf that separated from its 
menting it. Bison also defend themselves aggressively mother became mixed up in the middle of a Bison herd 
when attacked by Wolves (Canis lupus) (Carbyn and southwest of the Lamar Picnic Area. During the early 
Trottier 1988; Carbyn et al. 1993; MacNulty 2002), stages of this observation, filmmaker Ray Paunovich 
although Bison have been reported to be indifferent to saw a Bison knock the calf down and then repeatedly 
close, but non-attacking, Wolves (Catlin 1876-1: 254 butt it to the ground. 
cited by McHugh 1938; Goodwin 1939: 369 cited by RM watched a group of Bison encircle the calf, then 
McHugh 1938; and Soper 1941: 403; Carbyn et al. lick and sniff it. The calf got up and tried to walk off, 
1993). but several Bison chased it. One Bison butted the calf in 
We also have observed the same types of behavior of _ the side and knocked it down. Twelve additional Bison 
Bison toward wolves both when being attacked and came over to sniff it. The calf got up and ran off, but 
not being attacked in Yellowstone National Park. How- was chased and knocked down again by the Bison. 
ever, we have observed other types of Bison aggres- As long as the calf remained down or stood still, the 
_ siveness toward Elk and toward Wolves that has not — Bison just sniffed and licked it, but whenever it ran off, 
_ been reported, and we have seen another possible case they chased it and knocked it down. This sequence 
_ of Bison killing an Elk calf. Because our observations __ took place at least four times. A cow Bison later butted 
| considerably extend what is known about Bison aggres- _ the Elk calf several times as it lay on the ground. Then 
siveness and because observation of any ungulates chas- _ the herd walked away. The calf was motionless as the 
| ing predators are rare (Berger 1979), we report them Bison moved off and was possibly dead from all the 
below and propose an explanation for the possible adap- _ butting. RM then left, but Paunovich thought he saw 
tiveness of the aggressiveness. some movement from the calf after RM left. Whether 
__ Both Elk and Bison have inhabited the park for many the calf survived is unknown. 
_ decades, but Wolves were reintroduced to the ecosys- 2. 6 June 2001 — Bison Displace Wolves from Kill 


Bea an P73) and 1996 (Bangs et al. 1998). Wolves in "at 0529 hrs, RM spotted a fresh adult Elk kill with 


| Yellowstone prey primarily oo Elk (Mech et al. 2001) four yearling Druid Peak pack Wolves next to it south- 
_ but do kill Bison as well (Smith et al. 2000). RM made east of the Lamar Valley Picnic Area. The carcass was 


{ a peo auons in ns northeastem En of Yellowstone intact and the Wolves must have just killed it. Three 
ational Park through binoculars and spotting scopes Wolves were about to start feeding. 


_ with 60x power and recorded them by tape recorder: A Bison bull walked up to the carcass and sniffed its 


_ LDM independently observed and recorded with pad head. The Wolves went to the rear of the carcass and 
and pencil the observation on 20 March 2002. DWS 


HS 


116 


started tugging on that section as the bull continued to 
sniff the head area. This seemed to be the Wolves’ first 
feeding on the kill. Three more Bison bulls approached 
the carcass. One Wolf at the carcass walked off a few 
steps with a tucked tail, then came right back and 
snarled at the nearest bull. The bulls walked toward 
the three Wolves, and the Wolves backed off. Two of 
the bulls sniffed the Elk’s head. A Wolf sneaked back 
in and fed on the rear end. 

Soon six bulls were at the site. They had a stand-off 
with the three Wolves at 0537 hrs. When one Wolf 
tried to approach the carcass, a bull ran at it, and the 
Wolf backed off. The other bulls walked toward the rest 
of the Wolves, and they scattered. The six bulls kept the 
three Wolves away from the carcass through 0554 hrs. 
At that time, the bulls left the site, and the Wolves 
moved to the carcass and fed. The three Wolves were 
still feeding on the kill when RM left at 0615 hrs. 

In Wood Buffalo National Park, Alberta, mixed 
herds of Bison were observed forcing wolves away from 
wolf-killed Bison calves about 1 and about 4 months 
old (L. N. Carbyn, personal communication). 


3. 3 March 2002 — Bison Attack Wounded Elk Calf 

This incident was observed and video-taped by cine- 
matographer Shane Moore in the Blacktail area of 
Yellowstone National Park. The tape was viewed by 
RM who transcribed the details summarized below. 

The incident began when a herd of Bison ran 
0.15 km downhill to a 9-month-old Elk calf with blood 
on its neck. Several Coyotes (Canis latrans) were near- 
by. The Bison herd gathered near the Elk, and over a 
4-hr period, the Bison sniffed, licked, chased, and but- 
ted the Elk as the animal tried to seek refuge from the 
Coyotes by running into the Bison herd. The Bison but- 
ted the calf throughout the episode on 21 occasions 
during 40 minutes of videotaping, usually in the rear 
or side, and once knocked it into the air. At times the 
Elk was bedded or lying on its side when the Bison 
butted it. 

Partway through the video, it was apparent that the 
Elk’s abdomen was wounded, and an organ was hang- 
ing out. Moore believed that wound resulted from when 
he saw a Bison charge the Elk at full speed and hit it 
very hard in the side. Although the rest of the Bison 
herd then blocked the view, the calf stayed down for 
20 minutes and when it arose, its abdomen had been 
ripped open. 

Toward the end of the four-hour observation, the 
Bison drifted away from the wounded and bedded calf. 
As the calf continued to weaken, two Coyotes attacked 
it. Three additional Coyotes joined the attack, and the 
five Coyotes soon killed the calf. 


4. 20 March 2002 — Bison Displace Wolves from Kill 

Eleven members of the Druid Peak pack (1 1-months 
old and 23-months old; no adults) had been chasing 
Elk in the Hellroaring Creek area of Yellowknife Na- 
tional Park. At about 0931 hrs, the Wolves targeted a 
single cow Elk and pulled her down. A herd of about 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


39 Bison rushed to the downed Elk, chased the Wolves 
off, and surrounded the Elk tightly (0933 hrs). Sud- 
denly the Elk leaped up. Two Wolves approached the 
Elk and nipped at it. The Elk ran through the Bison 
herd, and the two Wolves pursued it. After about 30 m 
the Elk fell. The Wolves were on it again for about 30 sec 
when the Bison chased them off again (0936 hrs). Later 
the Elk arose and almost immediately collapsed. The 
Elk arose a third time, ran a short distance and col- 
lapsed. She later tried to stand again but could only 
get up on her hind legs. A Bison butted the Elk’s rear 
end and knocked her down, and the Bison herd 
surrounded the Elk again and kept the Wolves away. 
Each time a Wolf approached, a Bison would chase it 
15-30 m. After a few minutes, the Wolves headed 
away up a hill and bedded (0946 hrs). 

The Elk, while surrounded by Bison, kept raising her 
head and trying to get up. Usually the Bison stood 
closely around her but sometimes jumped back a few 
meters. The Bison remained tightly around the Elk 
from about 0939 hrs on. By about 1135 hrs, the Elk’s 
head was no longer up and her body lay flat; she seemed 
dead. Ravens landed on or next to her, although the 
Bison still surrounded her closely. Bison licked or 
sniffed (we could not determine which) the carcass 
intermittently for several minutes. 

The Wolves had returned at 0955 hrs and hung 
around 15-60 m from the Bison herd, and at various 
times some tried to reach the downed Elk. Each time, 
one or more Bison would skirmish with them and try 
to drive them off. 

About 1253 hrs, the Bison started moving away from 
the Elk, and by 1305 hrs Wolves moved in. However, 
the Bison quickly returned and ran the Wolves off. This 
happened several times when Bison were both east 
and west of the Elk. Eventually, however, the Bison all 
grouped east of the Elk, and the 11 Wolves began to 
feed. The Bison then charged the Wolves and ran them 
off. Then the Wolves returned and chased the Bison 
away. In the next few minutes there were two addition- 
al standoffs at the carcass, and both times the Bison 
chased away the Wolves. 

Such skirmishes continued as Wolves and Bison 
surged toward the Elk and each other, but gradually 
the Bison began to head a few meters farther east, and 
the Wolves became bolder. By 1313 hrs, the Wolves 
controlled the carcass and remained there and fed, while 
the Bison drifted off eastward. By 1350 hrs most of 
Wolves were done feeding and slept on a hill above the 
carcass, although individual Wolves fed later as well. 


5. 22 March 2002 — Bison Roust Sleeping Wolves 

In the same general area as observation 4 was made, 
eight of the 11 Druid Peak Pack Wolves seen on 20 
March were sleeping at 0909 hrs when six Bison lead- 
ing a larger herd (probably the same 39 seen there two 
days earlier) approached them. The Wolves arose, 
moved 100 m, and lay down again. Two minutes later, 
the Bison approached them to within 3 m, and the 


2004 


Wolves again moved off 100 m and lay down. At 
0918 hrs, a Bison approached one of the Wolves, 
which then confronted it; the other Wolves joined in 
harassing the Bison for 30 seconds. 

The Wolves then left at 0920 hrs and moved 90° and 
a few hundred meters and lay down on a rocky ridge. 
Eventually the Bison followed and rousted the Wolves 
from there. Several more times the Bison followed 
the Wolves and rousted them out of their beds. 


6. 16 April 2002 — Bison Bulls Displace Wolves from 
Kall 

At 0826 hrs, RM saw 11 Druid pack Wolves (the 
breeding male, two yearlings, and eight 1 1-month-old 
pups) chase Elk about 1.6 km west of the Lamar Ran- 
ger Station. They pulled down a cow Elk at 0858 hrs. 
A few Bison bulls started moving toward them from 
the north. Two Bison ran into the site and chased the 
Wolves off. A yearling and two pups came right back 
to the carcass. One of the bulls returned to the site and 
chased off one of the Wolves. 

At 0900 hrs several Wolves were feeding as four 
Bison bulls stood near the carcass. Two bulls charged 
the Wolves and drove them off, then stood close to 
the carcass. 

At that point, five Wolves watched the carcass from 
35 m away. The two bulls charged the breeding male 
and another Wolf, which fled eastward. The other 
Wolves ran to the kill and fed. The breeding male cir- 
cled around and joined them. 

A bull then walked to the carcass, and the feeding 
Wolves backed off a meter or so. The other nearby 
Bison approached the site. That bull charged the 
Wolves, and they ran off but came right back. 

Both bulls then stood next to the Elk carcass and 
sniffed it. The bigger bull charged the Wolves, but as 
before the Wolves soon stopped, and the breeding male 
and three others returned to the site. 

The two additional Bison bulls came toward the car- 
cass from the west. One of the bulls shook his head, 
and some Wolves backed off a step or two. Other 
Wolves stood their ground and faced the bull which 
was about 6 m away. The bull shook his head again, 
but the Wolves continued to feed and ignored him. 

The breeding male Wolf stepped away and ate some 
snow. When he returned to the carcass, a bull chased 
him off. The Wolf came right back. The same bull, 
which was the largest and most aggressive of the four 
bulls there, then drove all the Wolves off. The breed- 
ing male Wolf returned, and the big bull charged him. 
A second bull was standing a meter or so from the 
carcass. The big bull chased the breeding male Wolf 
southward, then returned to the site. The two bulls 
kept the Wolves from the kill until the bulls walked a 
short distance away at 09:12; the Wolves immedi- 
ately ran in. All 11 fed or were near the site. As the two 
bulls walked off to the north a pup followed them. 

Around 0920 hrs, three bulls approached the car- 
cass from the northeast as the Wolves continued to 


MEcH, MCINTYRE, AND SMITH: UNUSUAL BEHAVIOR BY BISON 


Ki, 


feed. One pup headed westward, but the others stood 
their ground. The bulls turned back before they reached 


the site. A fourth bull chased a Wolf to the north of the 
carcass. One bull then ran in toward the carcass, but the 
Wolves countercharged, drove him away, and resumed 
feeding. A fifth bull was north of the carcass but did 


not yet approach it. 

A sixth bull approached the kill, and a pup circled 
around to his rear and followed him. Another pup joined 
the first pup behind the bull. Two more of the original 
bulls approached the carcass from the north. The lead 
bull walked by the north side of the carcass with a 
raised tail. As another bull approached the site, the 
seven feeding Wolves backed off. At 0927 hrs three 
bulls remained at the carcass keeping the Wolves away. 
The breeding male Wolf stood nearby. He and a pup 
left, and the breeding male bedded north of the site at 
0929 hrs. Eight other Wolves stood near the carcass. 

At 0930 hrs two bulls started sparring with each 
other and began to drift away from the carcass. The 
breeding male Wolf and a yearling walked to the kill 
and fed. In the next minute other Wolves joined them. 
All six bulls remained away from the site, and the 
Wolves fed unmolested. 


Discussion 

The aggressiveness shown by Bison toward Elk and 
toward non-attacking Wolves is puzzling but might be 
some kind of hyperdefensive or hyperaggressive behav- 
ior. Predator harassment by several species of ungu- 
lates has been documented, and various explanations 
have been offered (summarized by Berger 1979), each 
involving ultimate reduction of predation risk. These 
explanations do not account for Bison attacking Elk, 
however. The most common element in our observa- 
tions and those reported earlier of Bison attacking Elk 
was the presence of a weak or downed Elk and the 
butting, sniffing, and mobbing of it. In the Elk inci- 
dents involving Wolves, perhaps the presence of the 
predators was only incidental and made the Bisons’ 
aggressiveness appear directed at keeping Wolves away 
from the carcasses or injured animals. 

In other words, our observations might involve two 
phases of hyperaggressive behavior: (1) a general 
aggressiveness toward various intruders, perhaps as 
generalized anti-predator behavior that causes the Bison 
to approach intruders, and (2) continued aggressive- 
ness toward intruders that do not flee, e.g., wounded 
animals or animals seeking refuge in the Bisons’ herd 
because these animals persist in remaining nearby. This 
would also include Wolves that are trying to feed on 
carcasses near the Bison herd. Bison are so large and 
powerful that when a herd acts aggressively, it can 
charge and attack any animal with impunity, includ- 
ing Wolves and Grizzly Bears (Ursus arctos). Thus it 
may be generally advantageous for them to advertise 
this ability by harassing intruders persistently without 
making fine distinctions as to type. 


118 


Acknowledgments 

This study was supported by the Biological Res- 
ources Division, U.S. Geological Survey, and the 
Yellowstone Center for Resources. 


Literature Cited 

Bangs, E. E., S. H. Fritts, J. A. Fontaine, D. W. Smith, K. 
M. Murphy, C. M. Mack, and C. C. Niemeyer. 1998. 
Status of gray wolf restoration in Montana, Idaho, and 
Wyoming. Wildlife Society Bulletin 26: 785-798. 

Berger, J. 1979. “Predator harassment” as a defensive strat- 
egy in ungulates. American Midland Naturalist 102: 197- 
199. 

Carbyn, L. N., and T. Trottier. 1988. Descriptions of wolf 
attacks on bison calves in Wood Buffalo National Park. 
Arctic 41: 297-302. 

Carbyn, L. N., S. M. Oosenbrug, and D. W. Anions. 1993. 
Wolves, bison and the dynamics related to the Peace-Atha- 
basca Delta in Canada’s Wood Buffalo National Park. 
Circumpolar Research Series (4). Canadian Circumpolar 
Institute, University of Alberta. 

Catlin, G. 1876. Illustrations of the manners, customs, and 
condition of the North American Indians. Two volumes. 
Chatto & Windus, London, U.K. 530 pages. 

Chapman, W., and L. Chapman. 1937. Wilderness wan- 
derers. Charles Scribner’s Sons, New York. 318 pages. 


THE CANADIAN FIELD-NATURALIST 


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Goodwin, G. C. 1939. The bison. Chapter XX in North 
American Big Game. Edited by Alfred Ely, H. E. Anthony, 
and R. R. M. Carpenter. Charles Scribner’s Sons, New 
York. 533 pages. 

MacNulty, D. R. 2002. The predatory sequence and the influ- 

ence of injury risk on hunting behavior in the wolf. Master’s 

thesis, University of Minnesota, St. Paul, Minnesota. 

Mahan, B. R. 1977. Harassment of an elk calf by bison. 

Canadian Field-Naturalist 91: 418-419. 

McHugh, T. 1958. Social behavior of the American buffalo 

(Bison bison bison). Zoologica 43: 1-41. 

Mech, L. D., D. W. Smith, K. M. Murphy, and D. R. 
MacNulty. 2001. Winter severity and wolf predation on a 
formerly wolf-free elk herd. Journal of Wildlife Manage- 
ment 65: 998-1003. 

Rush, W. M. 1942. Wild animals of the Rockies. Harper and 
Brothers, New York. 296 pages. 

Smith, D. W., L. D. Mech, M. Meagher, W. E. Clark, R. 
Jaffe, M. K. Phillips, and J. A. Mack. 2000. Wolf-bison 
interactions in Yellowstone National Park. Journal of Mam- 
malogy 81: 1128-1135. 

Soper, J. D. 1941. History, range and home life of the north- 
ern bison. Ecological Monographs 11: 347-412. 


Received 11 August 2003 
Accepted 4 June 2004 


Notes 


Morphology of Female Woodland Caribou, Rangifer tarandus caribou, 


in Saskatchewan 


W. JAMES RETTIE 


Department of Biology, University of Saskatchewan, 112 Science Place, Saskatoon, Saskatchewan S7N 5E2 Canada 
Present address: Northeast Science & Information Section, Ontario Ministry of Natural Resources, Ontario Government Complex, 
Highway 101 East, P.O. Bag 3020, South Porcupine, Ontario PON 1HO Canada 


Rettie, W. James. 2004. Morphology of female Woodland Caribou, Rangifer tarandus caribou, in Saskatchewan. Canadian Field- 


Naturalist 118(1): 119-121. 


I obtained morphological measurements from captured female Woodland Caribou in central Saskatchewan. I found that only 
girth was a good predictor of body mass in adult animals. I also determined that Woodland Caribou from Saskatchewan, though 
similar in size and mass to alpine Woodland Caribou in Yukon, are larger than migratory Woodland Caribou and smaller than 
forest dwelling Woodland Caribou from western Alberta and Yukon. 


Key Words: Woodland Caribou, Rangifer tarandus caribou, morphology, Saskatchewan. 


Body sizes and body masses of subspecies and pop- 
ulations of Rangifer tarandus (Eurasian Reindeer and 
North American Caribou) vary considerably (e.g., 
Dauphiné 1976; Thomas 1982; Reimers 1983; Chan- 
McLeod et al. 1995) in response to a wide range of cli- 
matic and other environmental conditions (Skogland 
1983; Reimers 1983). In North America, there is also 
a high degree of morphological variation within the 
Woodland Caribou subspecies (R. t. caribou). The 
best described populations of Woodland Caribou are 
those in Yukon (Gauthier and Farnell 1986; Kuzyk et 
al. 1999) and in Québec and Labrador (Parker 1981; 
Huot 1989). However, the subspecies has a more or 
less continuous distribution across northern Canada and 
the variations in climate, topography, and plant and 
animal communities across the range of the subspecies 
might be expected to favour different body sizes and 
masses in different areas. In particular, Skogland (1983) 
argued that morphology was density-dependent and 
related to the factors limiting population growth. Both 
Skogland (1983) and Chan-McLeod et al. (1999) 
reported that female R. tarandus that had reproduced 
successfully were significantly lighter the following 
winter than those that had not. Here, I present the first 
description of Woodland Caribou from central Saskat- 
chewan and a discussion of their morphology relative 
to limiting factors. 


Study Area and Methods 

As part of research into population dynamics (Rettie 
and Messier 1998) and behaviour (Rettie and Messier 
2000, 2001), I captured 43 female Woodland Caribou 
in central Saskatchewan (approximately 54° N to 
55°30’ N and 104° W to 109° W). All animals were 


captured in winter (dates between 13 December and 
14 March) between March 1992 and January 1995. 
At the time of capture I used a flexible steel tape to 
measure the following parameters from most animals: 
total body length, body length to base of tail, girth (all 
to the nearest cm), mandible, and metatarsal lengths (to 
the nearest 5 mm). Methods followed those of Dauphiné 
(1976), modified for application to live animals. I mea- 
sured body mass to the nearest kilogram for ten animals 
using an electronic load scale (Senstek, Saskatoon, 
Saskatchewan) suspended from a tripod or beneath a 
helicopter. I also extracted a tooth from most animals 
for aging. Further details on capture and aging appear 
in Rettie and Messier (1998). Animal capture and 
handling procedures followed animal care protocol 
number 920092 of the University of Saskatchewan. 

Using natural log transformed values from the eight 
adult animals for which I had complete sets of data as 
well as body mass, I applied a stepwise multiple linear 
regression to assess the relationship of body measure- 
ments to body mass (p = 0.05 to enter, p = 0.10 to 
exit) and then applied the resulting equation to the 
morphometric data for all animals. I then tested for 
differences in calculated body mass for adult females 
that were accompanied by a calf at time of capture and 
those that were without a calf. All statistical analyses 
were conducted using SPSS for Windows Version 10.0.7 
(SPSS Inc. 2000). 


Results and Discussion 

I had morphometric data from 34 adult animals 
(>31 months old at time of capture) and from three 
yearlings (19-22 months old at time of capture). The 
summary statistics for all measurements appear in 


119 


120 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE |: Morphological measurements and actual and calculated body mass of adult female Woodland Caribou in central 


Saskatchewan. 

Parameter n 
Actual body mass 9 
Calculated body mass” 32 
Body length to base of tail 33 
Total body length 33 
Metatarsal length 30 
Shoulder to hoof tip 15 
Girth 32 
Mandible length 32 


“ Body mass kg calculated using the equation arising from the regression analysis: e 


mean + | sd range 
131 + l6kg 96 -156kg 
LS eye like 96 —-160kg 
LOLS =p isiem 165 -—217cm 
200)=9 = lsicm 177. —230cm 
43.0 + 14cm 41.0 — 46.0 cm 
Il). ee W/m 102. —126cm 
AS) eee ikem 115) —154cm 
S14 ei ple8 cm 28.0 — 35.0cm 


1.76 In Girth — 3.78 


TABLE 2: Morphological measurements and actual body mass of yearling female Woodland Caribou in central 


Saskatchewan. 


Parameter n 


Actual body mass ] 
Body length to base of tail 3} 
Total body length 3 
Metatarsal length 3 
Girth 3 
Mandible length 3 


Tables 1 (adult animals) and 2 (yearling animals). 
When compared to Yukon Woodland Caribou, the 
animals in Saskatchewan are almost as tall at the shoul- 
der (115 cm vs. 116 cm), are the same girth (129 
cm), but are not as long (191 cm vs. 206 cm) as the 
alpine animals measured by Kuzyk et al. (1999). 
Saskatchewan caribou are smaller than forest 
dwelling Woodland Caribou from Yukon (Kuzyk et 
al. 1999) and from western Alberta (Gauthier and 
Farnell 1986) in all comparable parameters. 

From the regression analysis, I concluded that 
only girth was significantly related to body mass (p 
=0.04, r? = 0.78). I used the resulting equation, where 
girth is measured in cm: 


calculated body mass (kg) = e(!-76/Girth — 3.78 


to calculate body mass for all adult animals for 
which I had girth measurements. The calculated body 
mass values appear in Table 1. Figure 1 shows the 
relationship between girth and body mass and its 95% 
prediction interval. Untransformed data are presented 
in the figure while analyses were based on natural 
log transformed data. The wide prediction interval in 
Figure | suggests that further sampling is required to 
reduce uncertainty in the girth-body mass relationship. 
The relationship is particularly weak for smaller girths 
where the relationship is influenced by a single obser- 
vation. Though the summarised measurements present 
a reference point for future studies on morphological 
variation in Woodland Caribou, the prediction of body 
mass from body measurement and the equation pre- 


mean + | sd range 
98 kg 
GQ es. 3) con 170 —1175 cm 
185i lem 182 —1188 cm 
41.7 + 0.6cm 41.0 -— 142.0cm 
See eee Src 110 —1116cm 
28.2 + 2.0cm 26.0 — 130.0cm 


sented should be made cautiously (Cattet et al. 1997). 
Finally, I did not detect a difference in calculated body 
mass between adult animals with and without a calf-at- 
heel at time of capture (¢-test, t= 0.30, df = 30, p = 0.76). 

Skogland (1983) argued that body sizes in R. tar- 
andus were density-dependent, the larger body size 


200 body mass = @(1-76!n Girth - 3.78) 
*=0.78 

180 r S52 
P=0.004 - 

160 aa 


Body mass (kg) 
XD 
oO 


110 120 130 140 150 


Girth (cm) 


FiGurE 1. Relationship between girth (cm) and body mass (kg) 
for adult female Woodland Caribou in Saskatchewan. 
Solid line represents the equation derived through 
multiple linear regression; dashed lines represent the 
95% prediction interval of the equation. Filled squares 
represent data used to derive the equation. 


2004 


in North American Caribou being attributed to their 
low densities, a consequence of limitation by predation 
rather than by food resources. Amongst Woodland 
Caribou populations, the George River Caribou Herd 
in Quebec and Labrador is food limited and contains 
the smallest individuals (Parker 1981; Huot 1989). 
The body sizes of Woodland Caribou in Saskatchewan 
are larger than George River Herd animals and support 
the argument that Saskatchewan populations are not 
food limited (Rettie and Messier 1998). As with the 
larger animals in Caribou populations in western Al- 
berta and Yukon (Edmonds 1988; Kuzyk et al. 1999), 
predation is the likely proximate limiting factor for 
Saskatchewan Woodland Caribou populations (Rettie 
and Messier 1998). 

Skogland (1983) also suggested that predator in- 
duced constraints on lifetime reproductive success led 
to larger body size in North American Caribou by de- 
laying reproductive maturity in favour of increased 
somatic growth. However, yearling female Woodland 
Caribou in Saskatchewan were not only pregnant 
(Rettie and Messier 1998) but larger than both adult 
Reindeer (Skogland 1983) and adult migratory Wood- 
land Caribou (Huot 1989). Relative to observations in 
Norway, there does not appear to be a trade off being 
made between reproduction and somatic growth. In- 
stead, Saskatchewan Woodland Caribou appear able to 
mature as yearlings and still achieve large adult body 
sizes. My failure to detect differences in body mass 
between reproductive classes may represent a lack of 
difference, result from a poor predictive ability of my 
equation for body mass, or be a consequence of inter- 
annual variation as reported by Chan-McLeod et al. 
(1999). 


Literature Cited 

Cattet, M. R. L., S. N. Atkinson, S. C. Polischuk, and M. 
A. Ramsay. 1997. Predicting body mass in polar bears: 
is morphometry useful? Journal of Wildlife Management 
61: 1083-1090. 

Chan-McLeod, A. C., R. G. White, and D. E. Russell. 1995. 
Body mass and composition indices for female barren- 
ground caribou. Journal of Wildlife Management 59: 
278-291. 


NOTES 


Chan-McLeod, A. C., R. G. White, and D. E. Russell. 1999. 
Comparative body composition strategies of breeding and 
nonbreeding female caribou. Canadian Journal of Zoology 
77: 1901-1907. ; 

Dauphiné, T. C. 1976. Biology of the Kaminuriak population 
of barren-ground caribou. Part 4. Growth, reproduction 
and energy reserves. Canadian Wildlife Service Report 
Series Number 38. 

Edmonds, E. J. 1988. Population status, distribution, and 
movements of woodland caribou in west central Alberta. 
Canadian Journal of Zoology 66: 817-826. 

Gauthier, D. A., and R. F. Farnell. 1986. Comparison of 
caribou physical characteristics from Yukon and neigh- 
boring caribou herds. Rangifer Special Issue 1: 131-135. 

Huot, J. 1989. Body composition of the George River caribou 
(Rangifer tarandus caribou) in fall and late winter. Cana- 
dian Journal of Zoology 67: 103-107. 

Kuzyk, G. W., M. M. Dehn, and R. S. Farnell. 1999. Body- 
size comparisons of alpine- and forest-wintering wood- 
land caribou herds in the Yukon. Canadian Journal of 
Zoology 77: 1017-1024. 

Parker, G. R. 1981. Physical and reproductive characteristics 
of an expanding woodland caribou population (Rangifer 
tarandus caribou) in northern Labrador. Canadian Jour- 
nal of Zoology 59: 1929-1940. 

Reimers, E. 1983. Growth rates and body size differences in 
Rangifer, a study of causes and effects. Rangifer 3: 3-15. 

Rettie, W. J., and F. Messier. 1998. Dynamics of woodland 
caribou populations at the southern limit of their range in 
Saskatchewan. Canadian Journal of Zoology 76: 251-259. 

Rettie, W. J., and F. Messier. 2000. Hierarchical habitat 
selection by woodland caribou: its relationship to limit- 
ing factors. Ecography 23: 466-478. 

Rettie, W. J., and F. Messier. 2001. Range use and movement 
rates of woodland caribou in Saskatchewan. Canadian 
Journal of Zoology 79: 1933-1940. 

Skogland, T. 1983. The effects of density dependent res- 
ource limitation on size of wild reindeer. Oecologia 60: 
156-168. 

SPSS. 2000. SPSS for windows Version 10.0.7 SPSS Inc., 
Chicago. 

Thomas, D. C. 1982. The relationship between fertility and 
fat reserves of Peary caribou. Canadian Journal of Zoology 
60: 597-602. 


Received 3 December 2002 
Accepted 21 May 2004 


122 THE CANADIAN FIELD-NATURALIST Vol. 118 


Occurrence of Parasitoid Wasps, Baeus sp. and Gelis sp., in the Egg Sacs 
of the Wolf Spiders Pardosa moesta and Pardosa sternalis (Araneae, 
Lycosidae) in Southeastern Idaho 


LisA M. Coss! and VINCENT A. CoBB? 


Department of Biological Sciences, Idaho State University, Pocatello, Idaho 83209 USA 

Current Addresses: 

'Department of Biology, Cumberland University, Lebanon, Tennessee 37087 USA 

*Department of Biology, Middle Tennessee State University, Murfreesboro, Tennessee 37132 USA 


Cobb, Lisa M., and Vincent A. Cobb. 2004. Occurrence of parasitoid wasps, Baeus sp. and Gelis sp., in the egg sacs of the 
wolf spiders, Pardosa moesta and Pardosa sternalis (Araneae, Lycosidae) in southeastern Idaho. Canadian Field- 
Naturalist 118(1): 122-123. 


Egg sacs of the wolf spiders Pardosa moesta and Pardosa sternalis were sampled for two years during June to September in 
southeastern Idaho. Parasitoid wasps, Baeus sp. (Sceleonidae) and Gelis sp. (Ichneumonidae), were observed in the egg sacs 
of both Pardosa species. Of 322 egg sacs examined, 14.6% were parasitized. Parasitism of egg sacs occurred throughout 
most of the Pardosa egg sac-carrying season. 


Key Words: Wolf Spider, Pardosa moesta, P. sternalis, egg sac, predation, parasitic wasps, Baeus sp., Gelis sp., parasitoid, 


reproduction. 


Parasitic wasps are well known predators of spiders 
and may serve as either ectoparasites (e.g., externally 
feeding on the body) or endoparasites (e.g., feeding on 
eggs and developing young within egg sacs) (Foelix 
1982). Larval endoparasites that consume spiders eggs 
are classified as parasitoids (Roberts and Janovy 2000) 
with the most speciose family of parasitoid wasps be- 
ing the Ichneumonidae (Godfray 1994). To parasitize 
a spider egg sac, wasps insert their ovipositor into the 
wall of the egg sac and lay eggs which develop into 
larvae and predate the spider eggs (Austin 1985). 

Female wolf spiders (Lycosidae) have the conspic- 
uous behavior of carrying their egg sacs attached to 
their spinnerets. Although it would appear that such 
behavior could provide substantial maternal care, para- 
sitism of egg sacs still occurs. The wasps Baeus (Scel- 
ionidae) and Gelis (Ichneumonidae) have been found 
in previous studies to be common parasitoids of Lyco- 
sidae egg sacs (Kaston 1948; Edgar 1971a; Austin 
1985). 

Wolf spiders in the genus Pardosa are common 
ground-dwelling spiders throughout much of the north- 
ern hemisphere (Edgar 1971b; Dondale and Redner 
1990; Buddle et al. 2000). Their reproductive period 
may last several months (Edgar 1971b; Cobb 1992; 
Buddle 2000) and some species may produce multi- 
ple clutches during a single season (Edgar 1971b; Wolff 
1981). 

The main focus of this study was an investigation 
of the reproductive ecology of P. moesta (Banks) and 
P. sternalis (Thorell) in eastern Idaho. Cobb (1992) sug- 
gested that in southeastern Idaho both species have a 
tendency to produce multiple clutches within a season. 
During the study, P. moesta and P. sternalis females 
and their egg sacs were collected between one and five 
times per month from June through September of 1990 


and 1991. All collections occurred in a two hectare 
meadow near the base of Scout Mountain (Caribou Na- 
tional Forest, Bannock County, Idaho) at an elevation 
of approximately 2000 m. During the late spring and 
early summer the meadow was marshy, fed by a moun- 
tain stream. However, in late summer and early fall the 
meadow was dry and had been grazed by cattle. For 
each spider and corresponding egg sac we collected a 
variety of morphometric data, which included female 
mass, clutch mass, and clutch size. While examining 
the egg sacs we discovered several of them were para- 
sitized. 

Identifiable parasitoids of P. moesta and P. stenalis 
egg sacs were wasps of the families Scelionidae and 
Ichneumonidae. P. M. Marsh, of the USDA Agricultural 
Research Station, Beltsville, Maryland, identified the 
scelionid specimens as Baeus sp., and the ichneumonids 
at Gelis sp. All but two of the identifiable parasitoids 
for both Pardosa species were Baeus sp. Two egg sacs 
of P. sternalis were found with a single Gelis sp. 
each. The Gelis were collected on 23 June and 22 
July 1991. Pupal parasitoids inside the egg sacs could 
not be identified to species. Because not all individ- 
ual parasitoids could be identified, other parasitoid 
genera may have been present in the egg sacs. 

Of 134 egg sacs for P. moesta and 188 egg sacs for 
P. sternalis, 47 were infected with parasitoids (14.6% 
parasitism overall). Within these infected egg sacs 
281 parasitoids were counted. Parasitism rate was 
12.7% for P. moesta and 16% for P. sternalis (Table 
1). Egg sacs were found parasitized from June to Sep- 
tember for P. moesta and P. sternalis. During mid and 
late May 1990, a sample of egg sacs was collected from 
P. sternalis but none contained parasitoids. All of the 
P. sternalis egg sacs in the May collection contained 
eggs only and no spider instar stages, indicating egg 


| 


| 


2003 


TABLE |. Number of parasitized egg sacs for Pardosa moesta 
(n = 134) and Pardosa sternalis (n = 188) from southeastern 
Idaho. Data are represented as number of egg sacs with para- 
sitoids (number of egg sacs sampled). 


P. moesta P. sternalis 


Month 1990 199] 1990 199] 

June 0 (23) = 8 (37) Le CS) 
July 1 (40) 1 (9) 7 (45) Be (60) 
August 12 (25) = 6 (15) IP (G21) 
September 0 (27) 3(10) Dea alisia) Die (i) 


sac formation had begun only recently. The number 
of parasitoids per egg sac varied considerably. Mean 
(+ SE) number of parasitoids per egg sac for P. moes- 
ta was x = 5.1 + 1.4 (range = 2 - 15) in 1990 and 
x = 6.5 + 2.3 (range = 1 - 12) in 1991. For P. sternalis, 
the number of parasitoids per egg sac was x = 5.7 + 1.6 
(range = | - 37) in 1990 and x = 8.4 + 4.6 (range = 1 
- 35) in 1991. 

This study indicates that parasitism occurred through- 
out the egg carrying season and was highest in Aug- 
ust 1990 and September 1991. Interestingly, these high 
parasitism rates correspond to times when Pardosa in 
this region are producing smaller egg clutches (Cobb 
1992). Several species of Pardosa have their greatest 
reproductive output early in the reproductive season 
and produce smaller egg sacs and fewer eggs later in 
the season (Eason 1969). Potentially, these higher para- 
sitism rates in the late summer could have resulted in 
selection for greater spring reproduction in Pardosa. 
Although parasitism has been documented for several 
species of Pardosa (Eason et al. 1967), recorded levels 
of parasitism are uncommon and variable. Eason (1969) 
indicated egg parasitism rates of less than one percent 
in P. lapidicina in Arkansas, while Edgar (1971a) ob- 
served levels that ranged from 3% to 35% in Scotland. 
Our data indicate variable and sometimes high levels 
of parasitism as well. Such high levels of parasitism 
have been shown to reduce the number of spiders in 
the field (van Baarlen et al. 1994) as well as the re- 
cruitment of young (Edgar 1971a). Although we have 
documented seasonal variation in parasitism rate, we 
do not know if these Pardosa populations are being 
negatively impacted by parasitoids. 


Acknowledgments 

We thank P. M. Marsh of the U.S. Department of 
Agriculture for identifying parasitoid wasps and J. 
Redner of the Canadian Department of Agriculture for 
the initial identification of spider specimens. Funding 


NOTES 


for this project was provided partly by a grant to L. 
Cobb from the Idaho State University Graduate Res- 
earch Committee. 


Literature Cited 

Austin, A. D. 1985. The function of spider egg sacs in rela- 
tion to parasitoids and predators with special reference to 
the Australian fauna. Journal of Natural History 19: 359- 
370. 

Buddle, C. M. 2000. Life history of Pardosa moesta and Par- 
dosa mackenziana (Araneae: Lycosidae) in central Alberta, 
Canada. Journal of Arachnology 28: 319-328. 

Buddle, C. M., J. R. Spence, and D. W. Langor. 2000. 
Succession of boreal forest spider assemblages following 
wildfire and harvesting. Ecography 23: 424-436. 

Cobb, L. M. 1992. Reproductive tactics of two sympatric 
species of Pardosa (Araneae, Lycosidae). M.S. thesis. 
Idaho State University, Pocatello, Idaho. 59 pages. 

Dondale, C. D., and J. H. Redner. 1990. The insects and 
arachnids of Canada. Part 17. The wolf spiders, nursery- 
web spiders, and lynx spiders of Canada and Alaska 
(Araneae: Lycosidae, Pisauridae, and Oxyopidae). Agri- 
culture Canada Publication 1856. 

Eason, R. R. 1969. Life history and behavior of Pardosa 
lapidicina Emerton (Araneae: Lycosidae). Journal of the 
Kansas Entomological Society 42: 339-360. 

Eason, R. R., W. B. Peck, and W. H. Whitcomb. 1967. Notes 
on spider parasites, including a reference list. Journal of 
the Kansas Entomological Society 40: 422-434. 

Edgar, W. D. 1971a. Aspects of the ecology and energetics 
of the egg sac parasites of the wolf spider Pardosa lugu- 
bris (Walckenaer). Oecologia 7: 155-163. 

Edgar, W. D. 1971b. The life-cycle, abundance and seasonal 
movement of the wolf spider, Lycosa (Pardosa) lugubris, in 
central Scotland. Journal of Animal Ecology 40: 303-322. 

Foelix, R. F. 1982. Biology of spiders. Harvard University 
Press, Cambridge, Massachusetts. 

Godfray, H. C. J. 1994. Parasitoids: Behavioral and evolu- 
tionary ecology. Princeton University Press, Princeton, New 
Jersey. 

Kaston, B. J. 1948. Spiders of Connecticut. State Geology 
and Natural History Survey of Connecticut Bulletin 70. 
1020 pages. 

Roberts, L. S., and J. Janovy Jr. 2000. Gerald D. Schmidt 
and Larry S. Robert’s Foundations of Parasitology. 6" edi- 
tion. McGraw-Hill, Boston, Massachusetts. 

Van Baarlen, P. 1994. Eggsac parasitism of money spiders 
(Araneae: Linyphiidae) in cereals, with a simple method for 
estimating percentage parasitism of Erigone spp. eggsacs 
by Hymenoptera. Journal of Applied Entomology 118: 
217-223. 

Wolff, R. J. 1981. Wolf spiders of the genus Pardosa (Ara- 
neae: Lycosidae) in Michigan. Great Lakes Entomologist 
14: 63-68. 


Received 4 March 2002 
Accepted 14 June 2004 


124 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


The Northern True Katydid, Pterophylla camellifolia (Orthoptera: 
Pseudophyllidae), at Ottawa, Ontario 


STEPHEN J. DARBYSHIRE 


Agriculture and Agri-Food Canada, Central Experimental Farm, Saunders Building #49, Ottawa, Ontario K1A 0C6 Canada 


Darbyshire, Stephen J. 2004. The True Katydid (Prerophylla camellifolia) at Ottawa, Ontario. Canadian Field-Naturalist 


118(1): 124-126. 


Five males of the Northern True Katydid (Pterophylla camellifolia) are reported from Ottawa, Ontario (approximately 
45°25°N, 75°42’W), in 2001 and 2002 at three separate locations. Based on its distribution and habits extralimital 


occurrences in Ottawa are likely due to chance introduction. 


Key Words: Northern True Katydid, Pterophylla camellifolia, Ontario, distribution, insect dispersal. 


In mid-August 2001 two male Northern True Katy- 
dids, Pterophylla camellifolia (Fabricius), were heard 
calling at a site in an urban residential area of eastern 
Ottawa, Ontario (location 1, Figure 1). Calling contin- 
ued until about 9 October when daily minimum tem- 
peratures fell below freezing. Calling perches in trees 
were located between 5 and 10 metres above the ground. 
Several species of trees were utilized, including Sugar 
Maple (Acer sacharrum Marsh.), Norway Spruce 


(Abies balsamea (L.) Mill.) and Red Oak (Quercus 
rubra L.). Over the calling period they moved short 
distances (less than 50 metres horizontally) from one 
tree to another, but were never heard calling together in 
the same tree (it was assumed that the two males heard 
repeatedly were the same two individuals). Prerophylla 
camellifolia is not usually associated with coniferous 
trees and the Norway Spruce was occupied for only a 
single night. 


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FiGurE 1. Map of the Ottawa urban area showing the locations of calling males of Pterophylla camellifolia, approximately 
45°25°N, 75°42’ W. Location 1: two males in 2001; location 2: one male in 2002; location 3: two males in 2002. 


2004 


NOTES 


FIGURE 2. Male Pterophylla camellifolia from Ottawa, Ontario (location 3, Figure 1). Photographed 19 September 2002. 


Katydids were also heard calling in August, Septem- 
ber, and early October 2002 at two other locations 
within urban Ottawa (locations 2 and 3, Figure 1). At 
location 2 (Figure 1) a single male was heard calling 
from a large silver maple (Acer saccharinum L.) at a 
height of about 10 metres. It remained in the same tree 
throughout the observation period (about two months). 
At location 3 (Figure 1), two individuals were heard 
calling in a landscaped area around a large apartment 
complex. Perches were about three to four metres high 
in English Oak (Quercus robur L.) and Crab Apple 
(Malus sp.). Calling continued at these two locations 
until about 10 October, when night-time temperatures 
began to approach freezing. 

Since the insect is univoltine (Riley 1874; Caudell 
1906; Hebard 1941), searches were made of about 
1 km around location 1 in September 2002 and 2003 
and about 0.5 km? around locations 2 and 3 in Sep- 
tember 2003. Although repeated searches were made 
on warm evenings after complete darkness, no individ- 
uals were heard calling at location | in 2002, or at any 
of the sites in 2003. 

The range of Pterophylla camellifolia is reported to 
be from Massachusetts to north-central Florida, west- 
ward through southern New York, southern Ontario 
and Michigan to Iowa, Kansas, Oklahoma and eastern 
Texas (Hebard 1941; Vickery and Kevan 1983). Caul- 


_ field (1887) was the first to report the species in Ontario 


(under the name Platyphyllum concavum). The known 
distribution is primarily along the north shore of Lake 
Erie from the Niagara Peninsula to Essex County in 
southwestern Ontario and largely coincident to the 
Carolinian forest region (Vickery and Kevan 1985). 


The family Pseudophyllidae contains about 1000, 
mostly tropical, species. In Canada, the only represen- 
tative of the family is Prerophylla camellifolia. Although 
there are a few similar large green tettigoniid species 
in eastern Canada, P. camellifolia is readily distin- 
guished morphologically by its large size (about 25- 
50 mm) and its ovate, strongly convex tegmina (outer 
or front wings) which are slightly longer than the abdo- 
men (Vickery and Kevan 1983, 1985). The loud ono- 
matopoeic stridulations (chirps) also readily distinguish 
this species (audio file: Walker and Moore 2003*). 
Differences in pulse number and pulse frequency of 
the chirp occur throughout the range of the species 
(Alexander 1968) and with environmental conditions 
(Shaw 1975). The most common chirp is of three to 
five pulses followed by a short pause (Caudell 1906). 
Males heard in Ottawa produced three, or sometimes 
two or four, pulses per chirp. Hebard (1941) recognized 
two species of Prerophylla in the United States, one 
restricted to Florida and the other widespread in North 
America with considerable variation suggesting a sin- 
gle polymorphic species with five geographic races 
(subspecies). The northern type present in Ontario is 
P. camellifolia camellifolia. 

Blatchley (1920) stated that open forests are pre- 
ferred, although Hebard (1941) found the insect most 
common in dense forests, particularly where large oaks 
occur. Calling perches are usually in canopies when- 
ever tall trees are present, 25 to 100 feet (7.6-30.5 m) or 
above (Hebard 1941; Shaw and Carlson 1969). Where 
tall trees are absent, calling will occur from lower perch- 
es in small trees, orchards and shrubbery (Caudell 1906; 
Blatchley 1920; Hebard 1941). This is consistent with 
the observations of individuals in Ottawa. 


2003 


Throughout most of the species’ range males mature 
and begin calling in July and continue through to the 
advent of cold weather: late July to late September or 
early October in lowa (Shaw and Carlson 1969); late 
July to early October (rarely to early November) in 
Washington, D.C. (Caudell 1906); 30 July to 10 Octo- 
ber in Michigan (Cantrall 1968); and, 10 July to 27 
October in Indiana (Blatchley 1920). During the first 
severe frosts of autumn the insects fall to the ground 
(Hebard 1941) and perish unless the weather subse- 
quently moderates. 

Although possessed of tegmina and hind wings, 
Pterophylla camellifolia does not fly, the wings being 
used to parachute or glide from one perch to a lower 
one or to the ground (Caudell 1906; Hebard 1941; 
Shaw and Carlson 1969; Vickery and Kevan 1985). 
When on the ground or accessing higher perches they 
walk. Individuals normally travel only short distances 
during their lifetime (Caudell 1906; Hebard 1941), 
but may often move from tree to tree. 

The Northern True Katydid seems to be expanding 
its range northwards and westwards, with recent reports 
placing calling males in southern Minnesota (as far 
north as the Twin Cities area) (Tekiela 2002*), south- 
eastern North Dakota (Walker and Moore 2003*), and 
Colorado (Weissmann and Leatherman 1992: Walker 
and Moore 2003*). In Ontario recent sightings have 
been at Barrie (Sinclair 1998*) and Toronto (D. A. 
Sutherland, personal communication). 

What are the mechanisms by which this large flight- 
less insect could expand its range northward and west- 
ward to areas distant from its previous distribution? 
The new extralimital records in Ontario come from 
more-or-less urban regions, which suggests that inad- 
vertent human transportation is involved. The apparent- 
ly independent immigrations detected in the Ottawa 
area have been to well-established residential areas 
where commercial truck or rail traffic would not be a 
likely mechanism. It seems possible that nymphs or 
adults may be carried on non-commercial vehicles 
moving from southern regions during the early part 
of the summer. A trip from the contiguous range to the 
Ottawa area would take a minimum of 8-10 hours. 
The highway system and vehicle traffic have made such 
a trip possible for many decades now, yet immigra- 
tion has been noticed only recently. As it is one of the 
loudest insects in North America, the presence of 
mature males in residential areas is very conspicuous 
in spite of their cryptic green colouration and usually 
inaccessible calling perches. No females have been 
found in the Ottawa area yet and there is no evidence 
of an established breeding population. 


Acknowledgments 

D. A. Sutherland (Natural Heritage Information 
Centre, Peterborough, Ontario) provided observations 
on the occurrence of P. camellifolia in Toronto and 
discussed aspects of dispersal. Jean-Francois Landry 
and Henri Goulet (Agriculture and Agri-Food Canada, 


NOTES 


126 


Ottawa, Ontario) kindly read earlier versions of the 
manuscript and provided constructive criticism. 


Documents Cited (marked * in text) 

Sinclair, A. 1998. Pterophylla camellifolia “Northern True 
Katydid”. http://www.muskoka.com/~sinclair/katy.html 
[accessed 23 December 2003]. 

Tekiela, S. 2002. Minnesota Profile: Katydids. http://www. 
dnr.state.mn.us/volunteer/julaug00/katydids.html [accessed 
23 December 2003]. 

Walker, T. J., and T. E. Moore. 2003. Singing insects of 
North America. http://buzz.ifas.ufl.edu/index.htm [accessed 
23 December 2003]. 


Literature Cited 

Alexander, R. D. 1968. Arthropods. Pages 167-216 in Animal 
communication: techniques of study and results of research. 
Edited by T. A. Sebeok. Indiana University Press, Bloom- 
ington, Indiana. 686 pages. 

Blatchley, W. S. 1920. Orthoptera of northeastern America 
with especial reference to the fauna of Indiana and Florida. 
Nature Publishing Co., Indianapolis, Indiana. 784 pages. 

Cantrall, I. J. 1968. An annotated list of the Dermaptera, 
Dictyoptera, Phasmatoptera and Orthoptera of Michigan. 
The Michigan Entomologist 1: 299-346. 

Caudell, A. N. 1906. Class I, Hexapoda. Order XI, Orthop- 
tera. The Cryptophylli of the United States. Journal of 
the New York Entomological Society 14: 32-45. 

Caulfield, F. B. 1887. A sketch of Canadian Orthoptera. 
Eighteenth Annual Report of the Entomological Society 
of Ontario. Pages 59-72. 

Hebard, M. 1941. The group Pterophyllae as found in the 
United States (Orthoptera: Tettigoniidae, Transactions of 
the American Entomological Society 67: 197-219. 

Riley, C. V. 1874. Katydids. Pages 150-169 in Sixth annual 
report on the noxious, beneficial and other insects of the 
State of Missouri. 

Shaw, K. C. 1975. Environmentally-induced modification 
of the chirp length of males of the True Katydid, Ptero- 
phylla camellifolia (F.) (Orthoptera: Tettigoniidae). Annals 
of the Entomological Society of America 68: 245-250. 

Shaw, K. C., and O. V. Carlson. 1969. The true katydid, 
Pterophylla camellifolia (Fabricius) (Orthoptera: Tetti- 
goniidae) in Iowa: two populations which differ in behav- 
iour and morphology. Iowa State Journal of Science 44: 
193-200. 

Vickery, V. R., and D. K. McE. Kevan. 1983. A monograph 
of the Orthopteroid insects of Canada and adjacent regions. 
Lyman Entomological Museum and Research Laboratory 
memoir, number 13. 2 volumes, 1462 pages. 

Vickery, V. R., and D. K. McE. Kevan. 1985. The grass- 
hoppers, crickets, and related insects of Canada and adja- 
cent regions. Ulonata: Dermaptera, Cheleutoptera, Notop- 
tera, Dictuoptera, Grylloptera, and Orthoptera. The insects 
and arachnids of Canada. Part 14. Agriculture Canada, 
Research Branch Publication 1777. 918 pages. 

Weissmann, M. J., and D. A. Leatherman. 1992. Range 
extension of the northern true katydid, Prerophylla camel- 
lifolia (F.) (Orthoptera: Tettigoniidae Pseudophyllinae) into 
eastern Colorado. Journal of the Kansas Entomological 
Society 65: 448-449. 


Received 28 November 2002 
Accepted 6 January 2004 


2004 


L. Davip Mecu!? and SHAWN TRACY? 


Jamestown, North Dakota 58401-7317 USA 


127-129. 
} couver Island, British Columbia, Canada. 


Wolf (Canis lupus) densities are highly variable 
' (Mech 1970), depending generally on prey biomass 
| density (Keith 1983; Fuller 1989). Because prey bio- 
| mass density is related inversely to latitude, Wolves 
| tend to reach their highest densities at lower latitudes 
| (Mech and Boitani 2003). The highest reported natu- 
| rally occurring Wolf density is 14.1 Wolves/100 km? 
| for a pack of nine Wolves on Vancouver Island (Scott 
_ and Shackleton 1982). 

| Herein we report on a Minnesota Wolf pack with a 
| density higher for two years than the highest density 
| reported elsewhere. 


| Study area 

| We studied the Farm Lake Wolf pack (FLP) that in- 
| habited the Superior National Forest (SNF) of north- 
| eastern Minnesota where most of the Wolf’s diet is 
| White-tailed Deer (Odocoileus virginianus). The Wolf 
| packs in the SNF are spaced into territories (Mech 
| 1973, 1986) and the FLP territory, 10.0 km east of 
| Ely (48°N, 92°W), was surrounded by four or five other 
pack territories. The terrain, vegetation, and land use 
| in the territory are typical of the surrounding region 
| (Mech 1973). However, during winter, much of the ter- 
| ritory encompasses part of the Garden Lake deer yard 
(Nelson and Mech 1981). Deer densities there in the 
| late 1970s were estimated at 16-23 deer/km? (Nelson 
| and Mech 1981), and the deer population in the gen- 
eral area has increased considerably since then (Mech 
| and Nelson 2000). 


Methods 

‘| Wolves were live-trapped, radio-tagged, aerially 
|) radio-tracked weekly when possible, and aerially ob- 
|) served with their packs during winter. Estimates of FLP 
|) territory size were made for summer (1 April—30 Sep- 
ii tember), winter (1 October—31 March) and the entire 
|}, year (1 April-31 March) from 1 October 1997 through 
| 30 March 1999. 


NOTES 


‘Biological Resources Division, U.S. Geological Survey, Northern Prairie Wildlife Research Center, 8711 


127 


Record High Wolf, Canis lupus, Pack Density 


- 37" Street, SE, 


*University of Minnesota, Department of Fisheries and Wildlife, 1980 Hodson Hall, St. Paul, Minnesota 55108 USA 
| 3The Raptor Center, 1920 Fitch Avenue, University of Minnesota, St. Paul, Minnesota 55108 USA 


Mech, L. David, and Shawn Tracy. 2004. Record high Wolf, Canis lupus, pack density. Canadian Field Naturalist 118(1): 


_ This report documents a year-around Wolf (Canis lupus) density of 18.2/100 km? and a summer density of 30.8/100 km?, in 
a northeastern Minnesota Wolf pack. The previous record was a summer density of 14.1/100 km?, for a Wolf pack on Van- 


| Key Words: Wolf, Canis lupus, Minnesota, Superior National Forest, White-tailed Deer, Odocoileus virginianus. 


We used ArcView (copyrighted) GIS (ESRI Inc., 
Redlands, California) to estimate each period’s terri- 
tory area corresponding to the combination of all FLP 
radio-tagged Wolves. UTM coordinates of the FLP 
Wolf locations were imported into ArcView GIS and 
converted into point data. Minimum Convex Polygons 
(Mohr 1947) were then constructed using Animal 
Movement (Hooge and Eichenlaub 1997) extension 
to Arcview. 

By using observation curve analysis we determined 
when a territory was defined (Odum and Kuenzler 
1955). We subsampled period Wolf locations without 
replacement, increasing the number of samples (loca- 
tions) with each subsample treatment, and calculated 
territory area from each subsample. Five repetitions of 
each subsample were performed to generate a corres- 
ponding mean territory area for each subsample. The 
mean areas were then plotted against number of loca- 
tions in the subsample. We considered territories de- 
fined when there was <1% change in area (suggesting 
asymptotic behavior) as a result of the addition of one 
more sample. 

We determined pack size through the winter aerial 
surveys. The greatest number of Wolves seen was con- 
sidered the pack size. Vegetative cover prevented aerial 
surveys in summer. Pack sizes were divided by their 
territory size and then multiplied by 100 to give den- 
sity in Wolves/100 km’. 


Results 

We radio-tagged and tracked adult female Wolf 667 
and male pup 673 (born in 1997) from 1 October 1997 
through 30 March 1999 for this study (Table 1). A 
total of 127 Wolf locations were obtained, with one 
Wolf location defined as when one Wolf was at a point. 
If both Wolves were together, two locations were re- 
corded. Observation curve analysis suggested that 30 
locations were needed to accurately define the winter 
1997-1998 FLP territory, 50 samples for the 1998-1999 


128 


THE CANADIAN FIELD-NATURALIST 


TABLE |. Background data for calculations of Wolf density in the Farm Lake Pack, northeastern Minnesota. 


Period Number of Locations 
1 October 1997 — 30 March 1998 39 
1 April 1998 — 30 September 1998 44 
1 October 1998 — 30 March 1999 44 
1 April 1998 — 30 March 1999 88 


Vol. 118 
Number of Wolves Areakm? — Density/100 km? 
4 22.88 17.5 
— 19.50 30.8" 
6 32.88 18.2 
6 32.88 18.2 


“Assuming only six Wolves, the number observed during the following winter. 


territory, and 30 and 35 samples for the FLP summer 
and winter periods, respectively. In all these periods, 
these criteria were met (Table 1). 

During winter 1997-1998, the FLP contained four 
members and used an estimated area of 22.88 km?, a 
density of 17.5 Wolves/100 km. In winter 1998-1999, 
six FLP Wolves used an area of 32.88 km2, a density 
of 18.2 Wolves/100 km?, and in summer only 19.5 km? 
for a minimum density of 30.8 Wolves/100 km? 
(Table 1). 


Discussion 

The Wolf densities we found exceeded the highest 
previous record of 14.1 Wolves/100 km? for a Wolf 
pack on Vancouver Island (Scott and Shackleton 1982). 
Hebert et al. (1982), citing the Scott and Shackleton 
(1982) study, claimed a density of one Wolf per 6.3 
km, or 15.9/100 km?. However, Hebert et al. (1982) 
included a third pack not mentioned by Scott and 
Shackleton and presented no territory area data for this 
pack. Density estimates using radio-locations can be 
greatly influenced by the number and timing of loca- 
tions. Our locations were evenly distributed throughout 
the year and met the observation curve test (Odum and 
Kuenzler 1955), even though the number of locations 
available were fewer than recommended by others 
(Fritts and Mech 1981; Bekoff and Mech 1984; Ballard 
et al. 1987). No doubt the much smaller size of our ter- 
ritory explains why fewer locations were needed to 
define it. 

It is notable that the previous high record Wolf 
density (14.1 Wolves/100 km?) was based only on 
summer locations during a relatively short period (11 
April to 10 September). Furthermore, this period was 
before when most of the year’s mortality occurs (Fuller 
et al. 2003). Thus the earlier density is not strictly 
comparable to either our year-around or winter den- 
sities. 

The extraordinarily small FLP territory areas of 
1997-1998 and 1998-1999 resulted in the highest doc- 
umented Wolf density to date. This density was no 
doubt related to the high winter deer density in the area 
as well as to the relatively harsh winters during that 
period, with deep snow and extreme cold, that would 
have caused deer to concentrate with greater density. 
We know of no garbage dump or other regular source 
of food in this territory. 


Even so, our highest Wolf density (30.8 Wolves/ 
100 km?) occurred in summer, and it was a minimum 
estimate because it was based on the pack size the 
following winter, after any mortality may have oc- 
curred. Thus the winter deer density at first seems 
irrelevant. However, possibly such a high deer den- 
sity allowed the Wolves to add sufficient fat and to 
cache enough surplus food to help carry them through 
the summer. In any case, this study demonstrates the 
extreme density that Wolf densities can reach when 
prey is plentiful. 


Acknowledgments 

This study was supported by the Biological Res- 
ources Division of the U.S. Geological Survey and the 
U.S. Department of Agriculture North Central Forest 
Experiment Station. 


Literature Cited 

Ballard, W. B., J. S. Whitman, and C. L. Gardner. 1987. 
Ecology of an exploited Wolf population in south-central 
Alaska. Wildlife Monographs 98: 1-54. 

Bekoff, M., and L. D. Mech. 1984. Simulation analyses of 
space use: Home range estimates, variability, and sample 
size. Behavior Research Methods and Instrumentation 16: 
32-37. 

Fritts, S. H., and L. D. Mech. 1981. Dynamics, movements, 
and feeding ecology of a newly protected Wolf population 
in northwestern Minnesota. Wildlife Monographs 80: 1-79. 

Fuller, T. K. 1989. Population dynamics of Wolves in north- 
central Minnesota. Wildlife Monograph 105: 1-41. 

Fuller, T. K., L. D. Mech, and J. Fitts-Cochrane. 2003. 
Population dynamics. Pages 161-191 in Wolves: Ecology, 
Behavior, and Conservation. Edited by L. D. Mech and 
L. Boitani. University of Chicago Press, Chicago, Illinois. 

Hebert, D. M., J. Youds, R. Davies, H. Langin, D. Janz, 
and G. W. Smith. 1982. Preliminary investigations of the 
Vancouver Island Wolf (Canis lupus crassodon) prey 
relationships. Pages 54-70 in Wolves of the world. Edited 
by F. H. Harrington and P. C. Paquet. Noyes Publication, 
Park Ridge, New Jersey. 

Hooge, P. N., and B. Eichenlaub. 1997. Animal movement 
extension to Arcview. Ver. 1.1. Alaska Biological Science 
Center, U.S. Geological Survey, Anchorage, Alaska, USA. 

Keith, L. B. 1983. Population dynamics of Wolves. Pages 
66-77 in Wolves in Canada and Alaska. Edited by L. N. 
Carbyn. Canadian Wildlife Service Report Series (45). 

Mech, L. D. 1970. The Wolf: The ecology and behavior of 
an endangered species. Doubleday Publishing Company, 
New York. 


2004 


Mech, L. D. 1973. Wolf numbers in the Superior National 
Forest of Minnesota. U.S. Department of Agriculture 
Forest Service Research Paper NC-97. 

Mech, L. D. 1986. Wolf numbers and population trend in 
the Superior National Forest, 1967-1985. Research Paper 
NC-270. U.S. Department of Agriculture, Forest Service, 
North Central Forest Experiment Station, St. Paul, Min- 
nesota. 6 pages. 

Mech, L. D., and L. Boitani. 2003. Social ecology. Pages 
1-34 in Wolves: Ecology, Behavior, and Conservation. 
Edited by L. D. Mech and L. Boitani. University of 
Chicago Press, Chicago, Illinois. 

Mech, L. D., and M. E. Nelson. 2000. Do Wolves affect 
white-tailed buck harvest in Minnesota? Journal of Wildlife 
Management 64: 129-136. 


NOTES 


Mohr, C. O. 1947. Table of equivalent populations of North 
American small mammals. American Midland Naturalist 
37: 223-249. 

Nelson, M. E., and L. D. Mech. 1981. Deer social organ- 
ization and Wolf predation in northeastern Minnesota. 
Wildlife Monographs 77: 1-53. 

Odum, F. P., and E. J. Kuenzler. 1955. Measurement of 
territory and home range size in birds. Auk 72: 128-137. 

Scott, B. M. V., and D. M. Shackelton. 1982. A prelimi- 
nary study of the social organization of the Vancouver 
Island Wolf (Canis lupus crassodon). Pages 12-41 in 
Wolves of the world. Edited by F. H. Harrington and P. 
C. Paquet. Noyes Publication, Park Ridge, New Jersey. 


Received 25 August 2002 
Accepted 3 June 2004 


Extraordinary Size and Survival of American Black Duck, Anas 


rubripes, Broods 


JERRY R. LONGCORE and DANIEL G. MCAULEY 


United States Geological Survey, Patuxent Wildlife Research Center-Orono, 5768 South Annex A, Orono, Maine 04469- 


5768 USA 


Longcore, Jerry R., and Daniel G. McAuley. 2004. Extraordinary size and survival of American Black Duck, Anas 


rubripes, broods. Canadian Field-Naturalist 118(1): 129-131. 


Two female American Black Ducks (Anas rubripes) were initially observed during June 1982 with 20 Class Ib or 18-22 Class 
Ta-b ducklings in two wetlands in Hancock County, Cherryfield, Maine. Fifteen of 20 ducklings (75%) in one brood and 16 of 
18-22 ducklings (72-89%) in the other brood survived to fledge. These large broods probably resulted from post-hatch 
brood amalgamation. 


Key Words: American Black Duck, Anas rubripes, brood size, duckling survival, post-hatch brood amalgamation, Maine. 


Exceptionally large broods of North American dab- 
bling ducks (Anas spp.) that exceed average clutch 
size (8-10 eggs, Zammuto 1986) occur when females 
(1) lay extraordinarily large clutches, (2) lay eggs in 
nests of conspecifics (“pre-hatch brood amalgamation” 
(Pre-HBA), or brood parasitism; Eadie et al. 1988), 
or (3) hatch their own clutches and acquire the brood 
of another female (“‘post-hatch brood amalgamation” 
| (Post-HBA); Eadie et al. 1988). Pre-HBA, which can 
__ be either inter- or intra-specific, and post-HBA occur 
| infrequently in the Anatini; only 3 of 9 species of Ana- 
| tini were reported by Eadie et al. (1988) for either type. 
Tufts (1986) in Nova Scotia reported brood amalga- 
mation for three American Black Duck broods when 
_ he released orphaned ducklings near females with 
_ broods. Herein, we report two records of probable 
post-HBA resulting in two extremely large broods of 
wild American Black Ducks in Maine. 


_ Study Area and Methods 

| We observed the two broods in two Beaver (Castor 
| canadensis) flowages (Snake Flowage, 44°37'N, 68°06’; 
| BFA Flowage, 44° 39'N, 68°07'W) that were 20 km 
, northwest of Cherryfield, Maine, in township T10 SD, 
_ a forested area that has negligible acid-neutralizing 


capacity and low nutrients in wetlands (Norton 1980). 
We obtained morphometric and water chemistry char- 
acteristics of wetlands by methods of McAuley and 
Longcore (1988). We mapped and classified (Cowardin 
et al. 1979) both wetlands, and we sampled inverte- 
brates in one (Snake Flowage), as part of related field- 
work (J. R. Longcore, unpublished data). All obser- 
vations of broods followed the survey protocol of 
Longcore and Ringelman (1980). Morning visits on 
wetlands began 0.5 hour before sunrise and lasted two 
hours; the 2-h evening visit ended 20.5 hour after 
sunset. Broods were always sought on both wetlands 
simultaneously, and observers scanned wetlands with 
binoculars (7x50) and spotting scopes (20-60x) from 
elevated (4-5 m high) platforms. We backdated brood 
age (Gollop and Marshall 1954*) to determine approx- 
imate dates that first eggs were laid. 


Results 

During 3 June — 12 July 1982. we observed two 
American Black Duck broods that were twice (20 and 
18-22 ducklings) the size of average broods. Each 
brood was seen three times. Both broods were seen on 
the same wetland on the same day (8 June) by DGM. 
Both broods were observed on 12 July during the even- 


130 


Table 1. Characteristics of the two brood-rearing wetlands 
with the large American Black Duck broods in Cherryfield, 
Maine, 1982. 


Variable Snake Flowage BFA Flowage 

Wetland System, Palustrine, Palustrine, 
Class Forested Wetland Forested Wetland 

Basin area (ha) 4.9 4.5 

Surface water area (ha) 4.9 3.4 

% Submergents 10 78 

% Emergents 28 DD, 

% Flooded timber 4] 78 

pH (in situ) Sy) 6.13 

Alkalinity (ueq L"') 58.8 83.0 

Conductivity (uS cm!) 24.0 23.0 

Calcium (eq L"') 59.4 VS 

Phosphorus (ug L”') 70 170 

Color (Hazen units) 170 150 


ing visit, each by a different observer on a different 
wetland, thereby corroborating that two different broods 
existed as first identified by plumage characteristics 
(Gollop and Marshall 1954*). The brood of 20 dec- 
lined to 15 (75% survival) and the brood of 18-22 
declined to 16 (73 or 89% survival) at Class II size 
near fledging. The features of the two wetlands are 
described by the variables in Table 1. We sampled 
invertebrates in Snake Flowage, which contained the 
highest (P < 0.0001) number of aquatic Insecta per 
sample (mean = 257) compared with nine other wet- 
lands sampled in the area (J. R. Longcore, unpub- 
lished data). 


Discussion 

At Moosehorn National Wildlife Refuge, Maine, 
and at Lake Dalhousie, Nova Scotia (J. R. Longcore, 
unpublished data), we have observed Class II Amer- 
ican Black Duck broods of 10 ducklings, which are 
near mean clutch size. The two large broods, 
however, were twice the clutch size (mean +1 SD) of 
American Black Ducks in Maine (10.4 +1.3), in 
Vermont (9.6 +1.8; Coulter and Miller 1968), in 
Maryland (9.1 +1.8; Stotts and Davis 1960), or in 
Quebec (9.2 + 1.7; Reed 1970). Other studies of 
American Black Ducks have reported large clutches 
but rarely as large as the broods we observed. Reed 
(1970) reported sizes of mostly first clutches during 
12-25 April, as 13, 14, 15, and 17 eggs in Quebec. 
Both Coulter and Miller (1968) and Stotts and Davis 
(1960) reported clutch sizes of 14 or 15 eggs, and 
Krementz et al. (1991) reported that in 1982 
American Black Ducks nesting on islands in 
Chesapeake Bay averaged 10.2 + 3.0 eggs per clutch, 
ranging from 7-20 eggs, but it was unknown whether 
large clutches were from one female. 

For our two wetlands adequate food seemed avail- 
able for females to lay large clutches because the 
highest mean number of invertebrates per sweep net 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


sample was from Snake Flowage among 10 wetlands 
sampled (J. R. Longcore, unpublished data). By back- 
dating clutches we determined that these females, if 
they laid the entire clutch, wouid have initiated egg 
laying about 7 and 14 April, similar to early egg dates 
of 1-10 April for Maine and Vermont (Coulter and 
Miller 1968). Although ducks may adjust clutch size 
to environmental conditions (Skutch 1967), if these 
two females had laid 20 to 22 eggs per clutch, mass 
of the clutch would have been 1248 — 1373 g (62.4 g/ 
egg x 20 or 22 eggs), which would have equaled or 
exceeded mass of a female and been >2 times the 
average mass of a usual clutch (mean = 580.3 g, 
Arnold 1988). Therefore, it is possible that each female 
produced the entire clutch for these large broods, but 
it seems improbable. 

The possibility of intra-specific pre-HBA (hatch 
brood amalgamation) is supported by Stotts and Davis 
(1960) who reported two instances of American Black 
Duck females laying eggs in the nest of another fe- 
male. They (Stotts and Davis 1960: 145) also com- 
mented that “Eleven others [clutches] may have been 
the result of similar parasitism (a total prevalence of 
1.8 percent). An example of inter-specific pre-HBA 
also was recorded by Stotts and Davis (1960) who 
documented that an American Black Duck female 
began laying in a Mallard (Anas platyrhynchos) nest 
that contained five eggs. The Mallard laid four more 
eggs before deserting, but the American Black Duck 
laid 11 eggs to equal a 20-egg mixed clutch. Also, 
they reported that two American Black Duck females 
nested within 46 cm of each other on an offshore 
duck blind in Chesapeake Bay, Maryland; one female 
gradually incorporated the other’s clutch into her 
own and incubated all eggs (Stotts and Davis 1960: 
142). Large clutches associated with ducks nesting 
on islands, however, may be related to high nest 
density (i.e., 25.2 — 35.7 nests/ha on Bodkin Island, 
Chesapeake Bay, Krementz et al. 1991); but is uncom- 
mon elsewhere (e.g., 0.06 — 0.12 nest/ha in Maine 
bogs (Coulter and Miller 1968: 35). Because nest 
sites are not limiting in Maine or across the breeding 
range, nesting females are widely dispersed, except 
on islands (Coulter and Miller 1968), nests are well 
hidden (Bent 1923), and females are secretive when 
returning to nests in twilight hours (J. R. Longcore, 
unpublished data), it seems improbable that our large 
broods resulted from intra-specific pre-HBA (Beau- 
champ 1997). The third explanation for the large broods 
is intra-specific post-HBA. For this scenario, two addi- 
tional clutches of 10-12 eggs must have hatched at the 
same time and ducklings in those broods were then 
acquired by the females that we observed. We know 
that two other American Black Duck broods (i.e., of 
five and 10 ducklings) used Snake Flowage at the same 
time as the large broods. To account for this scenario 
of post-HBA, six American Black Duck broods of 
similar age must have been associated with these two 
wetlands. Concentration of broods on high quality 


2004 


wetlands is common (Longcore et al. 1998); indeed 
these two wetlands also supported one Green-winged 
Teal (Anas crecca), three Wood Duck (Aix sponsa), 
and four Hooded Merganser (Lophodytes cucullatus) 
broods. For the 20-duckling brood, JRL recorded on 
24 June that some ducklings appeared to be in different 
age Classes (i.e, I[b and IIc), which suggests post-HBA; 
however, the duckling brood of 18-22 appeared as all 
the same age to DGM. The two brood-rearing females 
were large, extremely attentive, and especially adept in 
eliciting rapid responses from ducklings by uttering a 
few low calls. Although females are capable of laying 
large clutches and the remote possibility of nest para- 
sitism (pre-HBA) exists, we conclude that the most 
plausible explanation for these two large broods was 
intra-specific post-HBA. 


Acknowledgments 

We thank P. E. Malicky for assistance during brood 
surveys and J. M. Eadie, T. W. Arnold, and A. J. 
Erskine for review and helpful suggestions on drafts 
of the manuscript. 


Documents Cited (* marked in text) 

Gollop, J. B., and W. H. Marshall. 1954. A guide for aging 
duck broods in the field. Mississippi Flyway Council 
Technical Report, Minneapolis, Minnesota. 14 pages. 


Literature Cited 

Arnold, T. W. 1988. Life histories of North American game 
birds: a reanalysis. Canadian Journal of Zoology 66: 1906- 
1912. 

Beauchamp, G. 1997. Determinants of intraspecific brood 
amalgamation in waterfowl. Auk 114: 11-21. 

Bent, A. C. 1923. Life histories of North American Wild 
Fowl. Order Anseres (Part). Smithsonian Institution, U.S. 
National Museum, Government Printing Office, Washing- 
ton, D.C. 

Coulter, M. W., and W. R. Miller. 1968. Nesting biology of 
Black Ducks and Mallards in northern New England. Ver- 
mont Fish and Game Department Bulletin 68-2. 74 pages. 

Cowardin, L. M., V. Carter, F. C. Golet, and E. T. LaRoe. 
1979. Classification of wetlands and deepwater habitats 
of the United States. U.S. Fish and Wildlife Service, FWS/ 
OBS-79/31. 103 pages. 


NOTES 13] 


Eadie, J. McA., F. P. Kehoe, and T. D. Nudds. 1988. Pre- 
hatch and post-hatch brood amalgamation in North Ameri- 
can Anatidae: a review of hypotheses. Canadian Journal 
of Zoology 6: 1709-1721. 

Krementz, D. G., V. D. Stotts, D. B. Stotts, J. E. Hines, and 
S. L. Funderburk. 1991. Historical changes in laying 
date, clutch size, and nest success of American black 
ducks. Journal of Wildlife Management 55: 462-466. 

Longcore, J. R., and J. R. Ringelman. 1980. Factors affect- 
ing waterfowl breeding density and productivity estimates 
in the northeast. Transactions of the Northeast Section, 
The Wildlife Society 37: 169-181. 

Longcore, J. R., D. A. Clugston, and D. G. McAuley. 1998. 
Brood size of sympatric American black ducks and mal- 
lards in Maine. Journal of Wildlife Management 62: 142- 
151. 

McAuley, D. G., and J. R. Longcore. 1988. Survival of juve- 
nile Ring-necked Ducks on wetlands of different pH. Jour- 
nal of Wildlife Management 52: 169-176. 

Norton, S. A. 1980. Geological factors controlling the sen- 
sitivity of aquatic ecosystems to acid precipitation. Pages 
521-531 in Atmospheric sulfur deposition, environmental 
impact and health effects: Proceedings of the Second 
Life Sciences Symposium, Potential Environmental and 
Health Consequences of Atmospheric Sulfur Deposition, 
Gatlinburg, TN, October 14-18, 1979. Edited by D. S. 
Shriner, C. R. Richmond, and S. E. Lindberg. Ann Arbor 
Science Publishing, Ann Arbor, Michigan. 

Reed, A. 1970. The breeding ecology of the Black Duck in 
the St. Lawrence estuary. Ph.D. dissertation, Laval Uni- 
versity, Quebec, Canada. 175 pages. 

Skutch, A. F. 1967. Adaptive limitation of the reproductive 
rate of birds. Ibis 109: 579-599. 

Stotts, V. D., and D. E. Davis. 1960. The Black Duck in the 
Chesapeake Bay of Maryland: breeding behavior and 
biology. Chesapeake Science 1: 127-154. 

Tufts, R. W. 1986. Birds of Nova Scotia. Third Edition. 
Nimbus Publishing and Nova Scotia Museum, Halifax, 
Nova Scotia. 

Zammuto, R. M. 1986. Life histories of birds: clutch size, 
longevity, and body mass among North American game 
birds. Canadian Journal of Zoology 64: 2739-2749. 


Received 28 February 2002 
Accepted 15 June 2004 


132 THE CANADIAN FIELD-NATURALIST Vol. 118 


Observations of Interactions between Puma, Puma concolor, and 
Introduced European Red Deer, Cervus elaphus, in Patagonia 


WERNER T. FLUECK 


CONICET (Consejo Nacional de Investigaciones Cientificas y Tecnoldégicas), Postal address: DeerLab, C.C. 176, 8400 
Bariloche, Argentina; e-mail: deerlab@infovia.com.ar 


Flueck, Werner T. 2004. Observations of interactions between Puma, Puma concolor, and introduced European Red Deer, 
Cervus elaphus, in Patagonia. Canadian Field-Naturalist 118(1): 132-134. 


Direct observations of interactions between native Puma (Puma concolor) and introduced European Red Deer (Cervus elaphus) 
in Patagonia are discussed with respect to the absence of evolutionary sympatry. Although the founding stock of European 
Red Deer had been lacking natural predation pressure for considerable time due to the previous extinction of large predators, 
these observations suggested that inherent antipredator behavior of European Red Deer toward this novel predator, once 
detected, was effective and may partially explain the success of European Red Deer as an invasive species. Puma behavior 
supported the view that they are a generalist predator which opportunistically utilizes new prey species like European Red Deer. 


Key Words: European Red Deer, Cervus elaphus, Puma, Puma concolor, introduced species, predation, behavior, interaction, 


Argentina. 


The role of large predators in regulating northern 
cervids continues to be debated. This function of large 
predators has to be viewed from a historical-evolu- 
tionary angle; how predators affect prey populations 
in modern human-modified settings can be expected to 
differ greatly (Berger 1998; Flueck 2000). The evolu- 
tion of large predators appears to be a systemic func- 
tional response as shown by re-evolving ecomorphs 
among very distant taxonomic groups. Furthermore, 
ecologically complete northern systems had several 
predator species with different hunting strategies and 
several prey species which thus prevented coevolution 
(Flueck 2000). Parts of Patagonia represent a natural 
experiment in south-temperate Argentina where Euro- 
pean Red Deer (Cervus elaphus elaphus) were intro- 
duced during the 1920s (Flueck and Smith-Flueck 
1993) to areas with native Puma (Puma concolor). As 
the Puma has only occurred in the Americas, these two 
species have no evolutionary history of sympatry. Fur- 
thermore, the native large predators had been extinct 
for a considerable time in areas of Europe where the 
European Red Deer originated, and immediate memory 
of such predators was thus absent. 

Hornocker’s (1970) work showed that direct obser- 
vations of attempts by Puma to capture prey was near- 
ly impossible and very few reports have been made 
previously, mostly stories by hunters (see Robinette 
et al. 1959). A few additional accounts are based on 
interpretation of tracks in the snow (Robinette et al. 
1959). During many years of studying Puma, Seiden- 
sticker et al. (1973: 31) never observed Mule Deer 
(Odocoileus hemionus) or American Elk (Cervus ela- 
phus nelsoni) mobbing a Puma, and I found no refer- 
ence to a cervid fighting off a Puma successfully. Here 
I report on direct observations of interactions between 
Puma and European Red Deer. 


Study Area 

The study area is in the western foothill portion of the 
province of Neuquén, Argentina (40°58'S, 71°12'W). 
The topography is primarily mountainous with most 
features formed by glacial processes. The dominant 
climate is temperate with main precipitation occur- 
ring between April and September. There is an abrupt 
precipitational gradient from west to east due to the 
rain shadow effect of the Andes which results in a 
strongly defined vegetation structure and floristic com- 
position. The study site is between 900 and 1200 m 
elevation and represents the ecotone between forest 
and steppe. Patches of forests are characterized by 
“Nire” trees (Nothofagus antarctica) and “Ciprés” 
(Austrocedrus chilensis) at lower elevations and are re- 
placed by “Lenga” trees (Nothofagus pumilio) at high- 
er elevations. Forest patches at lower elevations alter- 
nate with wet grasslands (“mallines”) with abundant 
growth of herbaceous plants, whereas at high elevation 
they are replaced by grass-dominated steppe of “Coirén 
Amargo” (Stipa speciosa var. major) and “Coirén Dul- 
ce” (Festuca pallescens), with variable occurrence of 
brush species like “Neneo” (Mulinum spinosum), 
“Calafate” (Berberis spp.) and “Espino Negro” (Colle- 
tia spinosissima). Riparian areas also contain galleries 
of small trees like “Radal” (Lomatia hirsuta), “Maitén” 
trees (Maytenus boaria) and “Laura” (Schinus pata- 
gonicus). 


Methods and Results 

Observations of interactions between European Red 
Deer and Puma were recorded while stalking alone 
during field work to collect deer for reproductive stud- 
ies. Here I report on two such incidents. 

On 10 November 1999 at 20:15 h I was returning 
to camp at a pace faster than stalking as it was getting 


2004 


late and the deer had not yet descended into the flat 
open feeding areas below surrounding hills which 
they use for cover during midday hours. The area I tra- 
versed was semi-open with patches of grassland inter- 
spersed with some patches of trees like Laura, Radal, 
and Cypress, typical of the acetone here. There were 
also patches of brush, predominantly Espino Negro 
and Calafate. As I was about to pass a stand of Radal, 
which was approximately 10 m in diameter, I heard the 
alarm bark of a European Red Deer cow about 20 m 
away coming from within the stand. I stopped imme- 
diately, believing that she must have seen me through 
the Radal trees, particularly as the same had happened 
only a short while earlier with a cow which was bed- 
ded in a similar tree patch. I thus waited in antici- 
pation that this animal might step out of the trees in 
curiosity, in which case I would have collected her as 
well. Looking with binoculars, I could see her move 
behind the trees and affirmed that she had not yet left. 

Suddenly, she barked again and came charging to- 
ward me through the patch of trees. Once she was in 
sight, I saw that she was pursuing an adult Puma, and 
it was clear that the barking was aimed at the predator 
and not at me. The Puma stopped shortly after get- 
ting out into the open, but as the cow came charging 
again, the Puma took off and the cow chased the cat 
around a circular patch of Espino Negro of about 7 m 
in diameter and | m in height. The fast chase took the 
animals around this bush patch twice, until the Puma 
apparently stopped out of sight behind the bush. The 
cow then stopped in front of it and commenced bark- 
ing again. Finally the cow trotted back toward the patch 
of trees on the same path where she had come from, 
only to turn around suddenly and charge toward the 
patch of bush, stamping in front of it and barking con- 
tinuously. Four times she repeated this behavioral pat- 
tern of trotting away and turning back, but when she 
finally reduced her aggressiveness and appeared about 
to retreat, I got ready to collect her. As she turned to 
enter the trees and was obviously walking away from 
the scene, I shot, which caused her to run through the 
tree patch on the same path on which she had origi- 
nally come out. 

After picking up my gear, I came around a bush 
and went towards the trees. Immediately, I saw the 
Puma jump out of a tree behind that patch of bush, 
and follow the deer on the same path through the trees. 
I then entered the tree patch, having to hunch down 
to pass through the trail. As I reached the other side, 
still in a hunched position, I saw the head of a Puma 
appear at about 15 m from behind a patch of bunch 
grass; it must have heard me coming through the trees. 
Looking through the binoculars, I determined it was 
a cub of the previous year. Still looking through my 
binoculars, I saw the head of the mother Puma pop 
up, and without hesitating she approached me to with- 
in 10 m. Apparently this was the first she knew of my 
presence and she began to snarl, which could be 


NOTES 


133 


clearly heard. After 30 seconds of staring straight at me, 
she sat down, but kept looking directly at me and snarl- 
ing periodically. She then appeared to relax in the sense 
that she would start to look from side to side, blink- 
ing slowly, and she was no longer in an alert position. 

As the sun had set and I needed to collect infor- 
mation on the cow before dark, I decided after about 
5 minutes to move on ahead. I stood up which im- 
mediately put the Puma into alert mode. She stood 
up, started to snarl continuously and stared at me. As 
I made a first few steps towards her, it became clear 
that she would rather move back than hold her posi- 
tion, so I continued to move forward. She retreated as 
much as I would move forward. Approximately 5 m 
out into the open, I could again see the cub which 
immediately retreated into the brush upon seeing me. 
At 15 m from the trees I found the dead cow, while 
the female Puma stood 10 m away still snarling. As I 
started to make brisker movements, like taking off 
the backpack, she retreated farther away and out of 
sight behind the brush. The thorax of the European 
Red Deer had already been torn open by the Puma. 

The European Red Deer cow was of average body 
size for this region (Flueck, unpublished data), except 
girth (116 cm) which was 10% below average. She 
had absolutely no fat reserves (sternum, rump, omen- 
tum, kidney). This was probably due to extreme drought 
conditions during spring through fall of 1999. Body 
conditions in fall of 1999 were so low that only 56% 
of adult females conceived (Flueck 2001) as compared 
to 100% in previous years. This 14-year-old female 
had given birth to a calf 1-2 days earlier as evidenced 
by the still enlarged uterus and the large but hard 
udder. Furthermore, in the trees I found remains of a 
calf which had been partially scavenged at least a day 
earlier; cause of death could not be determined. 

The second observation occurred on 17 November 
1996, while I was stalking in an area of hilly grassland 
interspersed with brush. At 18:15 h, I heard a Euro- 
pean Red Deer cow bark on the other side of a canyon 
and shortly after, I observed her running nervously 
back and forth in a semi circle at a patch of Espino 
Negro brush, about 5x10 m in area and | m tall. 
Then a calf cried and approximately 5 minutes later 
cried again. It was obvious that the female had a prob- 
lem with the calf. Shortly after, | saw an animal body 
appear from behind the brush patch and enter it. A 
few minutes later, the female stopped barking and 
trotted away down the hill. I then went to the spot and 
as I slowly approached the brush about where the ani- 
mal had entered earlier, an adult Puma fled crossing 
the brush patch. Right at the edge was a dead calf 
which was barely eaten. The Puma had dragged it from 
some distance when crossing the brush patch. The calf, 
a male, had body measurements indicating it weighed 
11 to 12 kg (Flueck, unpublished data), and several 
features indicating it was only a few days old (first 
incisors were out only 4 mm; umbilical scab). 


134 


Discussion 

Puma in Patagonia are large predators and compar- 
able to or surpass the largest races known from the 
nearctic (Iriarte et al. 1990). Franklin et al. (1999) 
reported average weights for female and male Pata- 
gonian Puma as 47.5 and 75.8 kg, respectively. Female 
Puma are known to increase their kill rate several 
fold when accompanied by cubs and can be expected 
to increase their aggressiveness. It is thus of interest 
that a European Red Deer female at the energetically 
most stressful period right after birthing coupled with 
nutritional stress from the drought was able to chase a 
female Puma which had young, up a tree. It indicates 
that the defense behavior of European Red Deer 
towards a discovered novel predator like Puma was 
effective in preventing the death of the mother. In the 
first case, the cause of the earlier death of the calf is 
unknown, and as it is common for a cow to remain in 
the vicinity of a recently born calf after it dies, it ex- 
plains the aggressive behavior toward the Puma which 
likely could have prevented predation on the offspring. 
In the second case, the mother could not prevent pre- 
dation of her calf by Puma; however, the learning event 
might enhance the probability that a future offspring 
might survive. Innate antipredator behavior of intro- 
duced European Red Deer therefore, appears success- 
ful in assuring adequate numbers of offspring to 
survive against the native Puma hunting strategy. The 
loss of exposure to large predators for the founding 
stock in their original habitat did not prevent the estab- 
lishment of the founding populations in Patagonia, 
indicating that rapid learning to defend against Puma 
must have occurred. Other cervids had become naive 
to their large predators which became locally extinct, 
but adapted quickly once these predators reappeared. 
This was clearly shown for Roe Deer (Capreolus 
capreolus), when Lynx (Lynx lynx) were reintroduced 
(Breitenmoser and Haller 1993), and Moose (Alces 
alces) with offspring after reestablishment of Wolves 
(Cants lupus) and Bears (Ursus arctos) (Berger et al. 
2001). 

On a larger scale, it is also clear that Puma in Pata- 
gonia, as the major predator of introduced European 
Red Deer, are not able to prevent them from increas- 
ing numerically (Flueck et al. 2003). Furthermore, as 
Puma and deer occur across extensive landscapes with 
no human-related mortality factors for Puma, I pos- 
tulate that intrinsic mechanisms are responsible for 
preventing Puma populations from reaching densities 
where they would exert a control over the deer popu- 
lations (Flueck 2000). Puma prey frequently on Euro- 
pean Red Deer indicating their generalistic and oppor- 
tunistic feeding behavior. Although about 80% of the 
natural mortality of adult female deer was due to Puma, 
annual survival rate of these deer remained high at 
91% (Flueck et al. 2004). 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


These observations support existing studies that the 
Puma behaves as a generalist predator which can oppor- 
tunistically adapt to new prey species. Furthermore, 
they demonstrate that European Red Deer have an 
ample repertoire of innate antipredator behavior which 
may partially account for their success as an invasive 
species. 


Acknowledgments 

I am grateful for the cooperation received from var- 
ious landowners and the Administracion de Parques 
Nacionales de Argentina. Thanks also to J. Smith-Flueck 
and P. S. Gibson for useful comments on the manu- 
script. 


Literature Cited 

Berger J. 1998. Future prey: some consequences of the loss 
and restoration of large carnivores. Pages 80-100 in 
Behavioral Ecology and Conservation Biology. Edited by 
T. Caro. Oxford University Press, Oxford, England. 

Berger, J., J. E. Swenson, and I. L. Persson. 2001. Recol- 
onizing carnivores and naive prey conservation lessons 
from Pleistocene extinctions. Science 291: 1036-1039. 

Breitenmoser, U., and H. Haller. 1993. Patterns of predation 
by reintroduced European lynx in the Swiss Alps. Journal 
of Wildlife Management 57: 135-144. 

Flueck, W. T., and J. M. Smith-Flueck. 1993. Uber das in 
Argentinien angesiedelte Rotwild (Cervus elaphus, L., 
1758): Verbreitung und Tendenzen. Zeitschrift fiir Jag- 
dwissenschaft 39: 153-160. 

Flueck, W. T. 2000. Population regulation in large northern 
herbivores: evolution, thermodynamics, and large predators. 
European Journal of Wildlife Research 46: 139-166. 

Flueck, W. T. 2001. Pregnancy rates of introduced red deer in 
Patagonia, Argentina after a period of drought. Ecologia 
Austral (Argentina) 11: 17-24. 

Flueck W. T., J. M. Smith-Flueck, and C. M. Naumann. 
2003. The current distribution of red deer (Cervus elaphus) 
in southern Latin America. European Journal of Wildlife 
Research 49(2): 112-119. 

Flueck, W. T., J. M. Smith-Flueck, and N. A. Bonino. In press. 
A preliminary analysis of cause-specific and capture-relat- 
ed mortality, and survival of adult red deer in northwestern 
Patagonia. Ecologia Austral. 

Franklin, W. L., W. A. Johnson, R. J. Sarno, and J. A. 
Iriarte. 1999. Ecology of the Patagonia puma Felis con- 
color patagonica in southern Chile. Biological Conser- 
vation 90: 33-40. 

Hornocker, M. G. 1970. An analysis of mountain lion preda- 
tion upon mule deer and elk in the Idaho primitive area. 
Wildlife Monographs 21: 1-39. 

Iriarte J. A., W. L. Franklin, W. E. Johnson, and K. H. 
Redford. 1990. Biogeographic variation of food habits and 
body size of the American puma. Oecologia 85: 185-190. 

Robinette, W. L., J. S. Gashwiler, and O. W. Morris. 1959. 
Food habits of the cougar in Utah and Nevada. Journal of 
Wildlife Management 23: 261-273. 

Seidensticker, J. C., M. G. Hornocker, W. V. Wiles, and J. 
P. Messick. 1973. Mountain lion social organization in 
the Idaho primitive area. Wildlife Monographs 35: 1-60. 


Received | September 2000 
Accepted 4 June 2004 


2004 NOTES 135 


Premieres mentions de la Couleuvre mince, Thamnophis sauritus 
septentrionalis, au Québec 


JEAN-FRANCOIS DESROCHES! et RICHARD LAPARE* 


'Collége de Sherbrooke, Département des Techniques d’écologie appliquée, 475 du Parc, Sherbrooke (Québec), 
J1K 4K1 Canada 

2Ministére des Transports du Québec, Direction de Laval-Mille-fles, 1725 boulevard Le Corbusier, Laval 
(Québec), H7S 2K7 Canada 


Desroches, Jean-Francois, et Richard Laparé. 2004. Premiéres mentions de la Couleuvre mince, Thamnophis 
sauritus septentrionalis, au Québec. Canadian Field-Naturalist 118(1): 135-137. 


Deux Couleuvres minces, Thamnophis sauritus septentrionalis, ont été trouvées les 12 et 13 aoat 2003, dans le 
comté de Pontiac en Outaouais, au Québec. II s’agit des premieres mentions de cette espéce dans la province. 
Des détails concernant les spécimens trouvés et les habitats sont présentés, de méme qu’une breve discussion 
sur la présence de cette espece au Québec. Trois autres Couleuvres minces ont été trouvées en 2004, dans le 
méme secteur, le méme qu’une autre a I’ fle-du-Grand-Calumet, a environ 30 km au nord-ouest. 


Two Ribbonsnakes, Thamnophis sauritus septentrionalis, were found on 12 and 13 of August 2003, in Pontiac 
County, Outaouais region, in Québec. These are the first records of the species for the province. Details on the 
specimens and habitats in which they were found are presented, as well as a brief discussion on the presence 
of Ribbonsnakes in Québec. Three additional Ribbonsnakes were found in 2004, in the same area, and another 
one on Ile-du-Grand-Calumet, about 30 km northwest. 


Mots-Clés : Couleuvre mince, Thamnophis sauritus septentrionalis, aire de distribution, premieré mention, 
habitat, Québec. 
Key Words: Northern Ribbon Snake, Thamnophis sauritus septentrioncilus, distribution, first record, habitat, 


Québec. 


La Couleuvre mince (Thamnophis sauritus) est pré- 
sente dans une grande partie de l’est de 1’ Amérique 
du Nord, de la Floride jusqu’en Ontario (Conant et 
Collins 1998). Au Canada, elle est représentée par la 
sous-espéce T. s. septentrionalis, laquelle se rencontre 
dans le sud de l’Ontario et sous forme de population 
disjointe en Nouvelle-Ecosse (Cook 1984). Malgré 
sa répartition limitrophe au Québec, cette couleuvre 
n’avait jamais été rapportée dans cette province. 7! s. 
septentrionalis mesure au plus 96,5 cm a l’age adulte 
et ressemble a premiére vue a la Couleuvre rayée de 
Pest (Thamnophis sirtalis sirtalis), présentant comme 
celle-ci un corps foncé marqué de trois rayures longi- 
tudinales généralement jaunes. On distingue la Cou- 
leuvre mince de la Couleuvre rayée par la position de 
ses rayures latérales, situées sur les 3*°™° et 4°™° ran- 
gées d’écailles du corps, son corps plus élancé et sa 
queue comparativement plus longue, et l’écaille pale 
bien définie devant chaque ceil (Cook 1984). La colora- 
tion des flancs, sous les rayures latérales, est souvent 
marron. La Couleuvre mince est considérée menacée 
en Nouvelle-Ecosse et en situation préoccupante en 
Ontario (COSEPAC 2002*), tandis que la Couleuvre 
rayée est tres commune dans ces provinces tout comme 
au Québec. 

Les 12 et 13 aoat 2003, dans le cadre d’un travail 
d’inventaire faunique effectué par le Ministére des 
Transports du Québec, dans un secteur circonscrit du 


comté de Pontiac, en Outaouais, nous avons décou- 
vert deux spécimens de la Couleuvre mince. Dans les 
deux cas les couleuvres se trouvaient sur une route de 
sable et de gravier et ont été apercues a partir d’un 
véhicule en mouvement. Le temps était ensoleillé et 
chaud, avoisinant les 30° Celsius. 

Le premier spécimen de la Couleuvre mince a été 
observé le 12 aoat a 18h10, sur le chemin Bristol 
Mines, a 1,2 km a l’ouest de la jonction avec le che- 
min Pontiac Station (45°29'29"N; 76°20'12"O), a 
environ 2 km au nord de la riviére des Outaouais. Il 
s’agissait d’un adulte mort écrasé, mesurant 57,6 cm 
du museau au cloaque, pour une longueur totale (in- 
cluant la queue) de 80,9 cm. Malgré que la couleuvre 
semblait avoir été tuée plusieurs heures auparavant, il 
a été facile de l’identifier par la position des rayures 
latérales et la couleur marron des flancs. Le second 
individu a été trouvé le 13 aodt a 16h53, sur un 
chemin reliant les immenses tas de débris miniers a 
la riviére des Outaouais (45°28'38"N; 76°20'15"O), a 
environ | km au nord de cette riviére. C’était un sub- 
adulte mesurant 29,7 cm du museau au cloaque, pour 
une longueur totale (incluant la queue) de 45,5 cm. 
Ce spécimen était vivant et 4 découvert sur le che- 
min. Des photographies des deux Couleuvres minces 
ont été prises par les auteurs. Les spécimens ont été 
déposés au Musée canadien de la nature (CMNAR 
35485, 35486). 


136 


L’habitat ot furent trouvées les Couleuvres minces 
est majoritairement forestier avec présence de milieux 
humides. Dans le premier cas, il s’agit dune forét 
mélangée a dominance de feuillus, ou poussent Acer 
rubrum, Populus grandidentata, Populus tremuloides, 
Fraxinus sp. et Thuja occidentalis. En bordure nord 
du chemin se trouve un étang 4 Quenouilles (Typha 
sp.) dense d’environ 250 m de longueur par 20 m de 
largeur. Dans le second cas, il s’agit d’une forét hu- 
mide a dominance de feuillus. On y trouve Betula 
papyrifera, Prunus sp., Acer rubrum, Populus balsa- 
mifera, Larix laricina, Fraxinus sp. et Alnus incana 
ssp. rugosa. A 40 m au nord, de chaque cété du ch- 
emin, un marécage est adjacent a la route. On y retrou- 
ve des arbres morts en milieu inondé. Des recherches 
effectuées dans les habitats avoisinants n’ ont pas per- 
mis d’observer d’autres Couleuvres minces. Quatre 
Couleuvres rayées de l’est, soit deux vivantes et deux 
mortes, ont été observées sur la route durant I’ inven- 
taire. 

En 2004, trois (3) autres Couleuvres minces adultes 
ont pu étre observées dans le méme secteur, qui couvre 
18 km?, et une (1) autre sur I’ Ile-du-Grand-Calumet, 
a quelque 30 km au nord-ouest. La premiere a été 
trouvée le 9 juin en bordure d’un étang a quenouilles 
pres d’un chemin de terre (45°29'10"N; 76°20'56"0). 
La seconde a été observée le 14 juin dans une flaque 
d’eau en asséchement, bordée de saules (Salix sp.), a 
10 m de la forét (45°29'40"N; 76°22'00"0). La troisiéme 
a été vue le 12 juillet sur une hutte de Castors (Castor 
canadensis) (45°31'51"N; 76°18'43"0). Le spécimen 
de I’[le-du-Grand-Calumet a été trouvé le 11 aofit sure 
une hutte de Castors (45°41'12"N; 76°38'31"0). Des 
photographies des Couleuvres minces ont été prises 
par l’un des auteurs (JED). 

La présence de la Couleuvre mince au Québec n’est 
pas surprenante puisqu’on la retrouve en Ontario a 
proximité, de l’autre c6té de la riviére des Outaouais 
(Tobias et Evans 1979; Cook 1981; Oldham et Weller 
2000*). Il est plut6t curieux qu’elle n’ait pas été 
rapportée plus tot dans la province. II s’agit bien sir 
d’une région peu habitée et sous-inventoriée par les 
biologistes, mais plusieurs inventaires spécifiques y ont 
été réalisés notamment pour les tortues. Il apparait pos- 
sible que sa ressemblance avec la Couleuvre rayée de 
Vest, tres commune dans la région, ait contribué a la 
garder dans |’anonymat durant toute cette période. 

La Couleuvre mince devient donc la 38'*™° espéce 
de l’herpétofaune au Québec et la 8°™° en ce qui con- 
cerne les couleuvres. La liste des couleuvres du Qué- 
bec comportait sept (7) espéces depuis 1927, année ot 
la Couleuvre brune (Storeria dekayi) fut découverte a 
Montréal (voir les données détaillées dans : Bider et 
Matte 1991). Les additions récentes a l’herpétofaune 
québécoise concernent les autres groupes (anoures, 
urodéles et tortues). Ce sont la Salamandre sombre 
des montagnes (Desmognathus ochrophaeus), décou- 
verte au Québec en 1988 (Alvo et Bonin 2003), la 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Tortue musquée (Sternotherus odoratus) en 1989 
(Chabot et St-Hilaire 1991) et la Rainette faux-grillon 
boréale (Pseudacris maculata) en 1991 (Bider et Matte 
1994). L’addition de la Couleuvre mince a la liste des 
reptiles du Québec revét une importance particuliére 
puisqu’il s’agit sans aucun doute d’une espéece trés rare 
dans la province et qu’elle est déja considérée en situa- 
tion précaire en Ontario et menacée en Nouvelle-Ecosse. 
Des inventaires supplémentaires seront nécessaires afin 
de préciser sa répartition et son statut au Québec. Fi- 
nalement, cette couleuvre vient probablement clore 
la liste de ’herpétofaune du Québec, si l’on se fie a 
la répartition actuellement connue des espéces. 


Remerciements 

Nous tenons a remercier le Ministére des Transports 
du Québec, et plus particulierement M. Ahmed Kho- 
dari, chef de service du Service des inventaires et du 
Plan, Direction régionale de |’Outaouais, pour avoir 
permis la réalisation de l’inventaire faunique au cours 
duquel les mentions de Couleuvres minces ont pu étre 
faites. Nous remercions également MM. Daniel St- 
Hilaire et Michel Lalancette, de la Société de la faune 
et des parcs du Québec, pour le prét des cartes topo- 
graphiques. Les spécimens de 2004 ont été trouvés par 
Mathieu Ouellette, Josiane Bergeron, Louis-Philippe 
Gagnon et Benoit Rivard. 


Documents cités (identifiés par * dans le texte) 

COSEPAC. 2002. Comité sur la situation des espéces en péril 
au Canada. http://www.cosepac.gc.ca/index.htm. 

Oldham, M. J., et W. F. Weller. 2000. Ontario Herpe- 
tofaunal Atlas. Natural Heritage Information Centre, 
Ontario Ministry of Natural Resources. http://www. 
mnr.gov.on.ca/MNR/nhic/herps/ohs.html (a jour le 09- 
11-2002). 


Littérature citée 

Alvo, R., et J. Bonin. 2003. Rapport sur la situation de la 
Salamandre sombre des montagnes (Desmognathus och- 
rophaeus) au Québec. Société de la faune et des parcs du 
Québec, Direction du développement de la faune. Québec. 
32 pages. 

Bider, J. R., et S. Matte. (Compilé par). 1991. Atlas des 
amphibiens et reptiles du Québec 1988-1989-1990, ver- 
sion détaillée. Société d’histoire naturelle de la vallée du 
Saint-Laurent et ministére du Loisir, de la Chasse et de la 
Péche du Québec. Québec. 429 pages. 

Bider, J.-R., et S. Matte. 1994. Atlas des amphibiens et rep- 
tiles du Québec. Société d’ histoire naturelle de la vallée du 
Saint-Laurent et ministére de l’Environnement et de la 
Faune du Québec, Direction de la faune et des habitats. 
Québec. 106 pages. 

Chabot, J., et D. St-Hilaire. 1991. Premiére mention de la 
Tortue musquée, Sternotherus odoratus, au Québec. Cana- 
dian Field-Naturalist 105 : 411-412. 

Conant, R., et J. T. Collins. 1998. A Field Guide to Rep- 
tiles and Amphibians of Eastern and Central North Ameri- 
ca. Third edition, expanded. The Peterson Field Guide 
Series. Houghton Mifflin Company, Boston. 616 pages. 


2004 


Cook, F. R. 1981. Amphibians and reptiles of the Ottawa 
District. Trail & Landscape 15 : 75-109. 

Cook, F. R. 1984. Introduction aux Amphibiens et Reptiles du 
Canada. Musée national des sciences naturelles et Musées 
nationaux du Canada. Ottawa, Canada. 211 pages. 

Desroches, J.-F., et David Rodrigue. 2004. Amphibiens et 
reptiles du Québec et des Maritimes. Editions Michel 
Quintin, Waterloo, Québec. 288 pages. 


NOTES 


137 


Revisée 16 Aott 2004. 
Tobias, T. N., et R. E. Evans. 1979. Range extension of the 
Ribbon Snake in Ontario. Trail & Landscape 13 : 180-181. 


Re¢u 20 aout 2003 
Acceptée 27 fevrier 2004 


Book Reviews 


ZOOLOGY 


Amphibian Decline: An Integrated Analysis of Multiple Stressor Effects 


Edited by Greg Linder, Sherry K. Krest, Donald W. Sparling. 
2003. SETAC North America, 1010 North 12th Avenue, 
Pensacola, Florida 32501-3367 USA. xxi + 345 pages. 
U.S.$98 Cloth. 


This volume contains presentations at the Workshop 
on the Global Decline of Amphibian Populations 
sponsored jointly by the Society of Environmental 
Toxicology and Chemistry (SETAC) and The Johnson 
Foundation 1-23 August 2001 at Racine, Wisconsin. 
Fifteen papers are included as 10 chapters, one with 
A-B and another with A-E designations for no appar- 
ent reason. 

Canadian data and perspective on the problem of 
declines are not neglected. Of the 35 contributing 
authors listed, 31 are from the United States but four 
are Canadian: Christine Bishop (Canadian Wildlife 
Service, Delta, British Columbia), David Cunnington 
(“freelance”’, Victoria, British Columbia), Martin 
Ouellet (Redpath Museum, McGill University, Mon- 
treal, Quebec), Bruce D. Pauli (Canadian Wildlife Ser- 
vice, Hull, Quebec). Ouellet is one of nine coauthors 
of Chapter 6 “Biotic Factors in Amphibian Population 
Declines”, and Bishop, Cunnington, and Pauli are half 
of the six coauthors of Chapter 7 “Physical Habitat 
and Its Alteration: A Common Ground for Exposure 
to Environmental Stressors”. 

None of the individual chapters is a particularly easy 
read and their results or conclusions often are neither 
clear-cut nor universal in their application. However, 
collectively, they effectively and repeatedly bring out 
the complications in separating climatic, chemical pol- 


Birds of Africa: From Seabirds to Seed-eaters 


By Chris and Tilde Stuart. 1999. MIT Press, Five Cambridge 
Center, Cambridge, Massachusetts, USA. 176 pages. U.S. 
$29.95 Cloth. 


This is an attractive book that appears to be in- 
tended as a typical “coffee table” volume. It is large 
in size (22 x 28 cm). profusely illustrated and very 
well laid out, with between 2 and 9 illustrations on 
each two-page spread. The plates themselves range 
from full page to quite tiny (under 4 cm’). 

The text is presented in 12 chapters plus an intro- 
duction, species list, and index. Each chapter is devoted 
to a group of bird families with some feature in com- 
mon. Thus we have “Birds of the Oceans’, “Birds of 
Inland Waters and the Coastline”, “LBJS”, and so on. 
Chapters then deal with each family in turn (grouped 


lutant, and biotic pressures, many of which doubtlessly 
combine to produce documented or supposed losses 
or declines. The texts are heavily laden with flow and 
modelling in 18 figure diagrams, 18 tables, 5 matrices 
and even two “test boxes” to visually present concepts 
and consequences. Each chapter has its own biblio- 
graphy. 

The concluding summary chapter effectively stresses 
the continuing, unresolved nature of the problems, but, 
in an inevitable rather self-congratulatory manner, 
arrives at the conclusion that this workshop was vital 
to promoting a much-needed exchange of expertise on 
various aspects of real and potential amphibian dec- 
lines and that it has created a network for further 
discussion and exchange. Although one could argue 
that this was happening anyway, the workshop did 
usefully focus discussions and the book spreads the 
state of knowledge. It is useful both for what research 
has been done and for what future approaches might be 
productive. One can hardly disagree with the funda- 
mental conclusion that further sharing of information 
as it develops will be essential to adequately defining 
future measures. If, eventually, there ever will be effec- 
tive conservation of the world amphibian fauna it will 
depend on a better understanding of the causes and 
reality of the declines. 


FRANCIS R. COOK 


Canadian Museum of Nature, Ottawa, Ontario KOG 1RO 
Canada 


in the sequence of their common names), giving the 
number of species that occur in Africa together with a 
brief account of the family characteristics, and some- 
thing about a sampling of the species. Occasionally 
there are boxes which treat a particular topic in some 
detail; for example, “The Dilemma of Island Birds”. 
The main body of the text enlivens the chronicle of 
numbers and species descriptions by concentrating on 
interesting anecdotes and “gee whiz” facts about the 
species under discussion. The authors are African and 
obviously have had considerable experience with the 
continent’s bird life. On the whole their statements 
seem accurate, although I was unable to check some 
of them, but some are incorrect. For example, Great 
Skua does not breed in the Southern Hemisphere, Kelp 


138 


2004 


Gull is not an African endemic, and the Australian 
Ibis is not conspecific with the Sacred Ibis. 

For me this book has some rather irritating problems 
that in sum seriously detract from its interest. Overall 
there is a lack of cohesion, mainly resulting from the 
organization of the text. The arbitrary chapter group- 
ings inevitably run into families that do not quite fit 
the category, or fit into more than one. I would be in- 
clined to place gulls and terns in the coastal waterbird 
chapter, lots of the “Seed-eaters” are classic LBJ’s, 
and “Terrestrial Birds” results in a strange collection 
of species with Pittas following Ostrich. This is ac- 
centuated by the layout within the chapters, with (for 
example) “Avocets and Stilts” at the beginning of 
Chapter 3, “Oystercatchers” and “Plovers and lap- 
wings” in the middle separated by Pelicans, and the 
“Waders or shorebirds” at the end. True, the system- 
atic list at the end is in order, but this does little to 
balance the sense of randomness that one gets from 
the text itself. 

Probably the authors would point out approaches 
of this kind are not unusual for books of the genre: 
they are not, after all, basic reference texts. Unfortu- 
nately, attractive though the layout is, there are prob- 
lems with the plates as well. In many cases the tinier 


The Firefly Encyclopedia of Birds 


Edited by Christopher Perrins. 2003. Firefly Books Ltd., 
3680 Victoria Park Avenue, Toronto, Ontario M2H 3K1 
Canada. 656 pages. U.S.$59.95. 


Naturalists across the continent strive to ever in- 
crease their knowledge of local wildlife, and all love 
to hear about wildlife from other localities. Birdwat- 
chers, for example, can become very knowledgeable 
about their local species, and the zealous ones will 
even study species from other areas within the same 
country, continent or even from around the world. 
But the vast array of bird species is overwhelming, 
and it takes a serious encyclopedia to pull them all 
together. Luckily for us, The Firefly Encyclopedia of 
Birds edited by Christopher Perrins is just what we 
needed — a good collection of text and photos of the 
birds of the world all in one volume. Nicely present- 
ed with tons of sharp photos illustrating everything 
from specific behaviours to simply jaw-dropping “‘cool- 
ness” (including several underwater pictures such as 
common murres on page 280-281, common kingfisher 
on page 368, and American dipper on page 521), the 
book is backed up by solid text summarizing well the 
basic tenets of ornithology. Moreover, each chapter 


BooK REVIEWS 


eo) 


illustrations are difficult to see properly, many of the 
poses are poor, and the selection of subjects pictured 
leaves something to be desired. Opening the book at 
random I find pp. 130-1 show two different pictures 
of Arrow-marked Babblers, one of the Southern Pied 
Babbler, and one of a Wallcreeper. The latter only 
occurs as a vagrant in North Africa, and the plate 
shows little of this species’ remarkable wing pattern, 
so it simply looks like a gray bird. Africa does have a 
creeper species, and the text on these pages also in- 
cludes the broadbills (Eurylaimidae). These species 
are not illustrated at all. 

In summary, this book brings together a large num- 
ber of photographs of African birds, some of them 
very good, together with a compilation of interesting 
facts and anecdotes on African bird families. My 
quarrels with content are really irrelevant if this is all 
you are looking for, or if you find this book attractive 
enough in itself to buy. Those who desire a thorough 
overview of the African avifauna should look else- 
where. 


CLIVE E. GOODWIN 


1 Queen Street, Suite 405, Cobourg, Ontario K9A 1M8 
Canada 


focuses on a bird family (the large number of species 
precludes a species-by-species analysis), and is accom- 
panied by a “factfile”. This sidebar summary provides 
a general description of size and habits, as well as a 
broad distribution map and my favorite, a visual com- 
parison of bird size to a human. 

The book also details several interesting specific 
behaviors, such as pigeon homing (pages 294-295), or 
the nest parasitism of Cuckoos (pages 316-317), but 
nothing on the degredation of the tundra by Snow 
Geese, a topic of high importance and actuality in 
North America. It also discusses extinct birds such as 
the Elephant Bird (page 21), the Dodo (page 291), 
and a short paragraph on the Passenger Pigeon (page 
293). The book provides a fantastic introduction to 
birds from all around the world, is extremely well 
presented, very reasonably priced ($75.00 CAN, 
hardbound), and constitutes a worthy, if not essential, 
addition to the library of all bird enthusiasts. 


SERGE LARIVIERE 


226 Rang Bois-Joly, St-Apollinaire, Québec GOS 2E0, Canada 


140 


Birds of Belize 


By H. Lee Jones. 2004. University of Texas Press, P.O. Box 
7819, Austin, Texas 78713-7819 USA. x + 444 pages. 
U.S.$60.00 Cloth, U.S.$34.95 paperback. 


This first edition covers the 574 species of birds 
recorded in the old British colony of Honduras, and 
now called Belize. Virtually all the birds are shown 
in colour plates, with accompanying text and 234 
range maps. The book has a brief introduction to the 
biogeography and climate of Belize and a how-to- 
use chapter. This latter is very important because it 
explains how the author has chosen to organize the 
book and select the “common” names, as well as the 
names in Maya, Spanish and the aboriginal language. 
The range maps, at the back of the book, mostly cover 
the resident species and are large enough to be a 
useful guide. 

The text is precise and clear and contains much 
useful information. For example, the call of the North- 
ern Potoo is described in terms that should enable 
anyone to recognize it. This is the key characteristic 
in separating the Common from the Northern Potoo. 
Another good example is the description of the Double- 
toothed kites, explaining the “Distinctive puffy white 
undertail coverts that spread to the sides of the rump 
.... This is a very good field character and this 
description captures it well. 

The artwork is by Dana Gardiner, the same artist 
that illustrated the Birds of Costa Rica by Stiles and 
Skutch. I found his style a little stiff, giving the birds 
a flat appearance and missing something of their jizz. 
These new renderings, while similar to those in Birds 
of Costa Rica, are an improvement. The Wedge-billed 
Woodcreeper illustration more clearly depicts its re- 
markable tail. The Emerald Toucanet is the correct 
subspecies, Aulacorhynchus prasinus prasinus, show- 
ing a white throat. However, I did have difficulties 


Herpetology: Third Edition 


By F. Harvey Pough, Robin M. Andrews, John E. Cadle, 
Martha L. Crump, Alan H. Savitsky and Kentwood D. 
Wells. 2004. Pearson Prentice Hall, Upper Saddle River, 
New Jersey, USA. ix + 726 pages. U.S.$93. 


In the preface to this edition of Herpetology, the 
authors state, “understanding amphibians and reptiles 
as organisms requires a perspective that integrates 
their morphology, physiology, behavior, and ecology 
and places that information in a phylogenetic context.” 
The authors have done a commendable job at fulfill- 
ing this daunting task. 

The text book is divided into four parts and each 
part is subdivided into chapters. Part one (What are 
amphibians and reptiles?) explores the field of her- 
petology, the place of amphibians and reptiles in ver- 
tebrate evolution, systematics and diversity of amphib- 
ians and reptiles, and biogeography. Part two (How do 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


with some plates, but I think this relates more to the 
printing than the artwork. For example, the depiction 
of Cedar Waxwing is far too intense and the elaenias 
are much too dark. The depiction of the Violet Sab- 
rewing is deep blue. I have never seen one in the field 
that did not look violet (as described accurately in 
the text and depicted accurately in Birds of Costa 
Rica) under any light conditions. In particular I 
checked the depictions of Scaly-breasted Humming- 
bird (poorly done in Birds of Costa Rica) and it is 
better, but I still think a novice would have trouble in 
the field using the plate for identification purposes. 

The nomenclature is fairly standard. The author 
retains Rock Dove for Columba livia (instead of the 
new “Rock Pigeon” — thereby reducing confusion 
with the orange-brown Rock Pigeon of South Africa, 
Columba guinea). | had to use the scientific name of 
Phaethornis longirostris to be sure the author’s Long- 
billed Hermit was the same as Stiles and Skutch’s 
(Western) Long-tailed Hermit. I am still confused as to 
the status and distribution of Passerini’s and Cherrie’s 
Tanager (a recent split of Scarlet-rumped Tanager). 
Only Paserrini’s Tanager is listed for Belize, which I 
believe is correct. 

This guide is just under 6” x 9” x 1.25” (15 x 22 x 
3 cm) and is too large for the average pocket. Al- 
though I have some reservations, | still think this book 
will make a useful field guide, especially if you use 
the text in conjunction with the plates. This book rep- 
resents a good stride forward in literature for this 
section of the Americas. If it is to be used in the field 
I would be tempted to take along a Mexican guide as 


a second reference. 
Roy JOHN 


2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario, 
K1J 6K5 Canada 


they work?) has chapters on temperature, energetics, 
reproduction, locomotion and feeding. Part three (What 
do they do?) covers movements, communication, mat- 
ing systems, diets and species assemblages. The fourth 
part (What are their prospects for survival?) discuss- 
es conservation of amphibians and reptiles in a single 
chapter. 

How does this edition differ from previous editions? 
The second edition was published in 2001 and hence 
this edition includes many references to recent pub- 
lications. There is also a greater emphasis on phylo- 
genetic analyses, particularly in the early chapters on 
systematics. This edition also features colour photo- 
graphs and colour distribution maps for the families 
of amphibians and reptiles. The colour maps do not 
really add much, but the colour photographs are effec- 
tive. A chapter on biogeography is one of the biggest 


2004 


changes in content in this edition. The authors have 
also wisely split the chapter on reproduction into two 
chapters, one each for amphibians and reptiles. The 
section on conservation has also been expanded, re- 
flecting the growing concern over many species in 
both groups. 

It is hard to find much wrong with this exhaustive 
examination of the current state of herpetology. The 
one conspicuous absence is a glossary, something that 
would surely be important in any comprehensive text- 
book such as this. Overall, the combined talents of 


Lizards: Windows to the Evolution of Diversity 


By Eric R. Pianka and Laurie J. Vitt. 2003. University of Cali- 
fornia Press, 2120 Berkeley Way, Berkeley, California, 
USA. xii + 333 pages. U.S.$45 Cloth. 


Individually, each author of this volume has more 
than 30 years of lizard study in various areas of the 
world. These include the southwestern (both) and 
southeastern (Vitt) United States, as well as Africa and 
Australia (Pianka) and Central and South America 
(Vitt). Each has included a personalized capsule bio- 
graphy of his background and interests, each empha- 
sizing what first attracted him to lizards. 

This book is number 5 in the University of Califor- 
nia Press series Organisms and Environments. The 
introduction explains that it grew from a resolve by 
authors 10 years ago. At that time, while working on 
a third lizard ecology symposium volume, they envi- 
sioned production of a comprehensive semi-popular 
book on the group. In a review of the earlier work, 
the 1994 Lizard Ecology: Historical and Experimental 
Perspectives, Robert W. Murphy (1995. The Canadian 
Field-Naturalist 109(1): 135-136), while generally 
praising its standards of “highest academic acumen”, 
thought it “not necessarily well suited for bedside 
reading by the merely curious. And it will not make a 
good coffee table book as there are no high-gloss 
photographs”. 

The latter criticism has been particularly well res- 
ponded to in the new volume. It is not only coffee- 
table size in design (28.6 x 22.5 cm) but has the req- 
uisite high-gloss photographs scattered throughout. 
These, often emphasizing activity or displays, splen- 
didly illustrate the diversity of behaviour as well as 
form and pattern for a well-chosen variety of lizards. 
There are also a scattering of graphs, tables, diagrams, 
and maps to illustrate particular points. Included are 
evolutionary relationships between groups of lizards, 
biotic and abiotic factors affecting a lizard’s well- 
being, avenue of heat gain and loss for a diurnal bask- 
ing lizard, comparison of mean percent use of the 
seven most important prey categories by neotropical 
and desert lizards, prey size vs lizard size, independent 
evolution of body type in Jamaica and Puerto Rico, 
and population size for species of land iguanas on 
islands of the Caribbean, to choose a few at random. 


Book REVIEWS 14] 


six experts with diverse research interests is hard to 
beat. After two previous editions, the authors have 
resolved most of the inevitable errors and contradic- 
tions resulting from integrating the writing of so many 
people. It is hard to imagine a better single volume 
overview of these fascinating creatures. 


DAVID SEBURN 


Seburn Ecological Services, 920 Mussell Road, RR 1, Oxford 
Mills, Ontario KOG 1S0 Canada 


The introduction raises the question: “what good 
are lizards?” This is first summarily dismissed with a 
curt retort, emphasizing the authors’ abhorrence of 
anthropocentrism, “what good are people?”. Subse- 
quently, the instructive value of lizards in relation to 
the ecosystem is attributed to their multitude of forms 
and the variety of habitats they have successfully 
occupied. They can thus effectively serve as “model” 
organisms for broad understanding of ecology and 
the diversity of animal life, a theme also emphasized 
in the earlier symposium volumes. 

Following the introduction are three major parts 
with seven, six, and two sections: (1) Lizard Life- 
styles: Evolutionary history and phylogeny, Getting 
around in a complex world, Lizards as predators, Es- 
caping predators, Social behavior, Reproduction and 
life history, and Reflections of a real world; (2) Liz- 
ard Diversity: Iguanians, From Geckos to Blind Liz- 
ards, From Racerunners to Night Lizards, Skinks, From 
Girdled Lizards to Knob-scaled Lizards, Monsters and 
Dragons of the Lizard World; (3) Synthesis: Historical 
perspective, Lizards and humans. In the initial section 
it is pointed out that as a group “lizards” is para- 
phyletic due to excluding the snakes. The latter are 
omitted as they traditionally have been treated as a 
separate equal group, though now recognized in con- 
temporary clarification as just an offshot within vara- 
noid lizards (as shown in Figure 1.4, page 16). 

The major partitions are followed by a four-page 
Taxonomic Summary (with number of species in each 
family or subfamily (if it has been subdivided) listing 
all included genera). The species totals are given (by 
major subdivisions) as [guania 1340+; Scleroglossa: 
Gekkota 973+, Incertae sedis 151+; Autarchoglossa 
(excluding snakes but inclusive of Scincomorpha: 
Lacertoidea, Scincoidea: Anguimorpha [including 
Varoidea]) 1745+; or collectively over 4200. The book 
concludes with a three-page Glossary (abiotic to zygo- 
dactyly), an 18-page references section, and a nine- 
page index. 

Throughout, the book is a pleasant mix of formal 
and informal styles. Scientific discussions have shuffled 
among them separate blocks of personal observations, 
often field experiences of one or the other author. As 
stated (page 7) “Throughout this book, we continu- 


142 


ally return to questions we asked as children in an 
attempt to encourage readers to open their minds and 
to ask questions”. This responds splendidly to the ear- 
lier reviewer’s wish for a text “well suited for bed- 
side reading by the merely curious”. Unfortunately, 
the large coffee table format and the glossy reflective 
pages make it awkward to handle in bed and read by 
bedside light. But as an up-to date comprehensive 
survey of world lizards it has no equal. It provides 


Raptors of Western North America 


Brian K. Wheeler. Princeton University Press, 41 William 
Street, Princeton, New Jersey 08540-5237 USA. 544 pages, 
625 colour photographs, 56 maps, U.S.$49.50 Cloth 


This book is both a tour de force and a labour of 
love, and the immediate new standard for diurnal 
raptors in western North America. Raptors are known 
for their variable plumages, and standard field guides 
cannot cover the almost infinite permutations and com- 
binations; hence, erroneous identifications are legion 
among regular birders, and even among raptor enthu- 
siasts. No full-time museum, government or university 
employee has dared to undertake a book of this scope; 
it is all the more astounding that this hiatus was filled 
by a truck driver. More incredibly, Wheeler has devel- 
oped the skills to take the great majority of the pho- 
tographs himself. His experience and enthusiasm are 
evident throughout. He has reviewed the literature 
carefully and has been given access to unpublished 
information from graduate students and others; for 
example, numbers of Northern Goshawk and Bald 
Eagle territories are provided for each state. There are 
nearly two pages about the ten-year cycle of the Snow- 
shoe Hare and Northern Goshawk, absent from most 
other raptor books; goshawks peaked at Duluth, Min- 
nesota, during the irruption years of 1972, 1982, 1992 
and 2001, mostly adults fleeing the mixed forest fol- 
lowing each Snowshoe Hare crash. 

Excellent historical accounts include notably that for 
the California Condor. There are four excellent glos- 
saries: general, anatomy and feather, plumage and molt, 
and displays. The detailed descriptions of raptor plu- 
mages are unprecedented and unequalled. Fortunately, 
important points are highlighted in bold, making the 
tedious mass of description more user-friendly. 

Readers will be amazed by the remarkable variabil- 
ity of plumage in species such as the Ferruginous 
Hawk (25 photos), Rough-legged Hawk (39 photos), 
and Swainson’s Hawk (40 photos), and especially the 
Red-tailed Hawk; variants within the latter species are 
depicted in 82 photographs, 22 pages of verbal des- 
cription and six maps. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


great perspective of their global importance, a fact 
strictly Canadian naturalists, who have only six spe- 
cies belonging three families of non-snake lizards to 
observe in their entire country, have little first-hand 
chance to appreciate. 


FRANCIS R. COOK 


Canadian Museum of Nature, Ottawa, Ontario K1P 6P4 
Canada 


John Economidy has contributed clear, accurate 
range maps for 33 species, six for the Red-tailed Hawk 
alone. The maps are incredibly precise, but I noticed 
four minor errors for Saskatchewan species: the Big 
Muddy area of Saskatchewan is omitted from the 
Golden Eagle; the two southern year-round localities 
for Bald Eagle should be winter only; recent small 
southerly extensions of Osprey and Northern Goshawk 
ranges are not shown; the two races of the Merlin are 
not known to overlap between Saskatoon and Prince 
Albert. The maps also give no indication of range 
changes in the past. 

Since this veritable encyclopaedia of western raptors 
(an eastern counterpart became available simultane- 
ously) is too heavy for most backpacks and too large 
for most glove compartments, most of us will use it 
as a reference in our libraries. One hesitates to quibble 
about the first edition of such a superb book. Yet the 
title is misleading, since owls are not included; inclu- 
sion of the term “diurnal” would have made the title 
more accurate. The ground squirrel has not lost half 
its range in western Canada, as it has in Idaho (page 
521). Use of dieldrin, not DDT, was coincident with 
the early 1960s crash of the Merlin on the Canadian 
prairies (page 447). Use of “very uncommon” some- 
times contradicts his definitions in the introduction. I 
detected few errors, but “verses” in place of “versus” 
jarred me (page 199). Wheeler has been involved in 
two previous hawk identification books, each excel- 
lent, each with William S. Clark as co-author. The 
first was Hawks in the Peterson Field Guide series and 
the second, A Photographic Guide to North American 
Raptors. For most general birders, either would suffice, 
although many plumages are omitted from both of the 
smaller books. If you are a raptor aficionado, don’t 
let the cost of the new book deter you; the 603 colour 
photographs easily justify the price and the 33 colour 
maps are more detailed than anything previously 
available. 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N OJ8, 
Canada 


I 


2004 


Raptors of Eastern North America 


By B. Wheeler. 2002. Princeton University Press, 41 Williams 
Street, Princeton, New Jersey, USA. xv + 439 pages, 
U.S.$45.00 Cloth 


Let me begin by saying this is a wonderful book. It 
gives a very detailed account of all the 26 full species 
of hawks, eagles and falcons occurring in eastern North 
America. It is profusely illustrated with extremely good, 
frame-filling, crisply focused photographs. These range 
from nine photographs for the Short-tailed Hawk to 
an incredible 82 of the Red-tailed Hawk. The text, des- 
pite its somewhat clipped English, is very informative. 
All of the known and recognizable sub-species are 
included, both in the text and, where valid, in the 
photographs. The level of detail given in the plumage 
descriptions is far greater than in any other book I 
have read. The range maps are sized as appropriate to 
the species they cover. For widespread birds the map 
depicts all of eastern North America. However, larger 
scale maps are used for such localized birds as Flor- 
ida’s Snail Kite. 

This is easily the best photographic-style guide I 
have seen and is a real testament to the author’s dedi- 
cation. Thus it is the most thorough guide to this group 
of North American birds you can purchase. The full- 
color, superb photographs alone are worth the price 
of the book. The author has included all the plumages, 
races, and colour variations possible. They cover a 
bewildering range of plumages for species that can 
be annoyingly difficult to identify in the field. This 
makes it an almost essential book for serious bird- 
watchers and ornithologists and a great resource for 
beginners trying to grapple with the plumage variabil- 
ity. I do not consider this book to be a portable field 
guide, but a reference work. It is a little too large and 


Turtles and Tortoises 


By Vincenzo Ferri. 2002. Firefly Books Ltd., 3680 Victoria 
Park Avenue, Toronto, Ontario M2H 3K1 Canada. 255 
pages. $24.95 


This compact little book is a translation of the 1999 
Italian publication, Tutto Tartarughe e Testuggini. It 
can best be thought of as a photographic guide to 152 
species of turtles — roughly half of the species alive 
today. 

The book opens with an introduction covering the 
biology, evolution, classification and conservation of 
turtles. The bulk of the book is occupied by species 
accounts. The species accounts are grouped geograph- 
ically into oceans (sea turtles) and six terrestrial areas: 
Palearctic, Afrotropical, Oriental, Nearctic, Neotropical 
and Australian regions. There are 152 numbered spe- 
cies accounts and additional unnumbered accounts 
sprinkled throughout the book. It is unclear why some 
species accounts are not numbered, although the un- 
numbered accounts do not have a colour photo (they 


BOOK REVIEWS 143 


heavy (due to the thick, high-quality paper). More 
important, this would be too precious a book to risk 
damaging in the field. 

I searched through for errors and did not notice any. 
In fact, I was impressed by the thoroughness of the 
coverage. For example, the author has correctly includ- 
ed the spotty northern locations for Osprey in Lab- 
rador. 

However, I do have one major problem with this vol- 
ume. It is a companion volume to Raptors of Western 
North America, a slightly larger book covering 33 
species. Thus, there are 23 species common to each 
book. The species accounts and photograph are almost 
identical, except that the range maps are different. This 
means there is a good deal of unnecessary repetition. 
I can only assume that some marketing guru felt that 
two versions would sell better than one. Combining 
into one book would increase the size from 544 to 
about 600 pages to allow for the text for two addition- 
al species and the eastern range maps for all species 
(plus a little added to the index.). This would presum- 
ably increase the price to around U.S.$60. Each book 
is a worthwhile purchase on its own. If you buy the 
western book you will miss the accounts for Snail Kite 
only. If you buy the eastern guide you will lose the 
accounts for nine western raptors. If you buy both you 
will get more than 75% repetition, in essence wasting 
U.S.$30, to get the coverage of one extra species plus 
the relevant range maps. If you can afford only one 
book, buy the western guide. 


Roy JOHN 


2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario, 
K1J 6K5 Canada 


do have a colour illustration) or map. The species ac- 
counts are brief (some less than 100 words), with most 
ranging from half a page to a full page in length. Each 
account has standardized subheadings: Family, Distri- 
bution and habitat, and Characteristics. Some of the 
accounts also have a “Situation” subheading describing 
conservation issues. 

Perhaps because it is a translation, this book is 
plagued with errors. Translation-type errors include 
some unusual common names. For example, the Bog 
Turtle (Clemmys muhlenbergii; now Glyptemys muh- 
lenbergii) is called Muhlenberg’s Turtle. That may well 
be the English equivalent of the Italian common name, 
but that term is not used in North America. Similarly, 
the Painted Turtle (Chrysemys picta) is referred to as 
the Painted Tortoise. In addition, the genus is mis- 
spelled “Chrysemis.” Errors in content also abound. 
The author asserts that Eunotosaurus is the earliest 
known ancestor of turtles, yet this theory is no longer 


144 


widely accepted. It is also stated that some turtles can 
hibernate for 2-3 months, a gross understatement of 
the 6+ months that some turtles spend in hibernation. 

The species accounts are also fraught with errors 
or misleading information. For some North American 
species, the author lists individual U.S. states where 
the species is present, yet the list is rarely complete. 
The Blanding’s Turtle (Emydoidea blandingii) account 
omits any mention in the text or the map of the Nova 
Scotia population. The author accepts the Mississippi 
Map Turtle (Graptemys pseudogeographica kohni) as 
a species (G. kohni), even though the species account 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


suggests it is a subspecies. There are also some curi- 
ous omissions. There are six species accounts for mem- 
bers of the genus Graptemys yet the most wide-ranging 
species, the Northern Map Turtle (G. geographica) is 
not included. Overall, one has the sense that the text 
was assembled quickly, or by using outdated reference 
materials, and that it was not reviewed by a scientific 
expert. Enjoy the photos but don’t believe everything 
you read. 

DAvID SEBURN 
Seburn Ecological Services, 920 Mussell Road, RR 1, Oxford 
Mills, Ontario KOG 1SO Canada 


What Good are Bugs? Insects in the Web of Life 


By Gilbert Waldbauer. 2003. Harvard University Press, 79 
Garden Street, Cambridge, Massachusetts, USA. 384 pages. 
U.S'$29295 


Waldbauer has written several other popular books 
about insects, and these have been met with high 
acclaim; I have not read those tomes, and so see his 
most recent work with unbiased eyes. 

What good are bugs? focuses on the interactions 
insects have with other animals and plants, both living 
and dead. Several chapters in each category illustrate 
the myriad ways in which insects, wittingly or unwit- 
tingly, are key to earth’s ecosystems as we know them 
today. Topics range from seed dispersal to recycling 
dead animals and the control of animal and plant 
populations. Truly, the scope of this book is close to 
all-encompassing, and is written in a style that will 
not be condescending to the informed naturalist or 
biologist, but will be readily consumable by the bud- 
ding naturalist, too. 

There are a few factual problems, as well as some 
opinions that are more typical of the entomophobic 


component of the public, that were surprising and dis- 
appointing to see in a book which promotes insects. 
Several times Waldbauer has insects “attacking” plants, 
when he really means “feeding on” plants. No one 
would ever say that deer or rabbit attack plants, it’s 
no different with insects; they are just feeding on the 
plants, a point that naturalists, biologists, and surely 
this author should understand. “Attacking plants” is 
an expression used by certain people or industries to 
incite action, to justify eradication, to gain sympathy 
from the uninformed, and should itself be eradicated 
from our vocabulary. 

The book ends with a chapter-by-chapter listing of 
selected readings. I like this method of listing refer- 
ences since the reader can readily choose among 
works only in the topic of interest. Overall, this is a 
book well worth having. 


RANDY FE. LAUFF 


Department of Biology, St. Francis Xavier University, 
Antigonish, Nova Scotia B2G 2W5 Canada 


Belugas in the North Atlantic and the Russian Arctic 


Edited by M.P. Heide-Jorgensen, @.Wiig, and D. G. Pike. 
2002. NAMMCO Science Publication 4, The North At- 
lantic Marine Mammal Commission, Polar Continental 
Centre, N-9296, Tromso, Norway NOK 150. 270 pages. 


The greater part of this symposium is devoted to 
those beluga whales that migrate through Baffin Bay 
and Davis Strait between the eastern part of the 
Canadian Arctic and West Greenland, and are difficult 
to follow across the deep water (and impossible in 
the dark season). Moreover, it is a difficult publica- 
tion to review because it represents “work in progress’’, 
the results of which are not yet fully understood by 
the authors themselves. Three main techniques of study 
were (1) mitochondrial molecular genetics from tissue 
sampling of restrained or dead animals; (2) aerial pho- 
tographic surveys; (3) tracking individuals by means 
of satellite radio tags attached to implants in the dorsal 
ridge of temporarily trapped belugas. The results are 


too tentative to summarize easily. Many groups of bel- 
ugas observed in summer in arctic estuaries are essen- 
tially matrilines, that is, adult females with young ani- 
mals of both sexes, while most adult males may move 
independently in different ways. There is also the prac- 
tical concern that catches of belugas at southwest Green- 
land, where there is most open water and the largest 
catching boats, appear to exceed recruitment. The exact 
area from which these animals come is unknown. 

After this it is a pleasure to move on to simpler 
studies! An overall survey of Russian and Siberian 
arctic waters from the mainland was carried out in 
the course of ice studies for belugas, narwhals and 
Greenland or Bowhead whales, plus the few Grey 
whales that enter the Arctic Ocean. There is now little 
hunting for them in this half of the Arctic. 

Lastly, a summary is given of up-to-date knowledge 
of the numerical status of belugas that inhabit, year- 
round, the less than 200 linear km of the St. Lawrence 


2004 


estuary in Quebec. This is the most biologically pro- 
ductive part because of year-round vertical water mix- 
ing. Aerial photographic surveys of numbers are there- 
fore simple to carry out, but it is necessary to measure 
the percent of animals diving too deeply to be regis- 
tered visually or by camera form the air. M. C. S. 
Kingsley and team did this using a helicopter hover- 
ing at an altitude that did not disturb the animals. The 
correction factor turned out to be more than two, so 
that estimates of numbers needed to be more than 
doubled, giving a total of more than 1200. Moreover, 
a series of estimates made since even casual hunting 
ended, ca. 1979, up to the last recorded survey in 1998, 
showed that the population has been growing slowly. 

I add to this review some more recent information that 
several belugas, including one radio-tracked animal, 
moved in autumn 2002 from SE Hudson Bay to off 


The Firefly Encyclopedia of Insects and Spiders 


Edited by Christopher O’Toole. 2002. Firefly Books Ltd., 
Willowdale, Ontario 


I will first state my biases and say that I’m not a big 
fan of encyclopaedias. Having got that out of the way, 
I must say this book was engaging, excellently laid out, 
and an absolute joy to read. The photographs ranged 
from very good to excellent to amazing. The full-frame 
images of even small insects startled me with their 
clarity. Diagrams were liberally used, effective, and 
also of exceedingly high quality. Each chapter was 
written by one or more experts, and I enjoyed reading 
them all. The majority of insect orders are covered, 
as are most of the arachnids (not just the spiders as 
the title alludes). 

Although this book could not serve as a university 
text in entomology, if I were to offer a bug course at 
the high school level, I would use this book. I have 
seen many entomology texts, and none are as visu- 
ally appealing as this book. This book would not sub- 
stitute for a field guide because of its size, but the qual- 
ity of the text, illustrations and photos far exceeds any 
of the field guides I’ve used. 

Interspersed with the taxonomic accounts which 
dominate the book were photo stories, fact files and 
special features. These articles ranged from less than 
a page to two pages and provided a different tangent 
on insect and arachnid life. As with the rest of this 
book, these articles were well done. 


BooK REVIEWS 


145 


Nain, Labrador (D. W. Doidge, personal communi- 
cation), showing that therefore some animals “bleed” 
from the Arctic into the Labrador Current. These 
must be the source of infrequent but not rare animals 
which reach the east coast of Newfoundland (Curren 
and Lien. 1998. Canadian Field-Naturalist 112(1); 28- 
31). These were noted by Kingsley in this book, but 
have not been tracked further. Animals coming from 
the St. Lawrence estuary are also known to “bleed” 
into waters around the Maritime Provinces as far as 
the Bay of Fundy and even to New Jersey (various 
references). It therefore seems very unlikely that the 
St. Lawrence population is isolated genetically from 
the main Arctic population of belugas. 


D. E. SERGEANT 
Box 745, Hudson Heights, Quebec JOP 1JO Canada 


The weaknesses were very few. My underlying 
feeling was that I couldn’t be certain as to the in- 
tended audience. The wording is usually so straight- 
forward that children from eight or nine years old 
should be able to read it; however, technical ter- 
minology creeps in now and again, and would likely 
give those same early readers quite a pause. For the 
most part, the technical lingo is kept to a minimum, 
and is often isolated from the main text in‘ sidebars. 
This book is billed as an encyclopaedia, and the 
publishers I think were striving to market the book to 
a wide audience. 

My only quibble about the actual content was that 
the minor taxa were mentioned only in a list along- 
side the more dominant taxa, and not given their own 
words or photographs to describe them. I realize that 
no book that features the ricinuleids more than the 
beetles will ever sell, especially to the general public, 
but a photograph and a quarter page or so of text for 
each of these minor taxa would have been appropriate. 
Overall though, a great reference book, with won- 
derful visual appeal. 


RANDY F. LAUFF 


Department of Biology, St. Francis Xavier University, 
Antigonish, Nova Scotia B2G 2W5 Canada 


146 


BOTANY 


Biotic Forest Communities of Ontario 


By Norman Duncan Martin and Norma M. Martin. 2003. 
Commonwealth Research. Belleville, Ontario, Canada. 
195 pages. $10.00 paper. 


Martin and Martin set out to identify the basic for- 
est communities of Ontario, describe their biotic com- 
position, and consider the successional relationships 
among them. While they draw on an impressive amount 
of personal research and a broad literature survey, the 
book lacks focus and therefore may have a limited 
value to a general audience. 

A fundamental component of any scientific study 
is that the work must be repeatable. When presenting 
study results it is therefore essential that the methods 
are Clearly laid out. This is not the case here. The read- 
er is informed that the authors tallied vegetation in 
transects in various types of forest. A single map of 
transect locations is provided, with numerous exam- 
ples of data sheets. No details are provided to explain 
the selection of transect locations nor which aspects 
of the vegetation were sampled or how. There is also 
no explanation of the analysis itself, other than to 
acknowledge the influence of the “schools of inter- 
pretation” of Clements, Curtis, Hills, Whittaker and 
others. These were indeed important workers in this 
field, but listing their names doesn’t allow the reader 
to critically examine the results of the current study. 

The bulk of the book is devoted to descriptions 
and tables illustrating the ten forest community types 
identified by the authors. There are interesting obser- 
vations here, but the sheer quantity of poorly organized 
data makes it difficult to appreciate. Most of the data 
is presented as tables showing various measures of 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


abundance in selected samples. The only graph pre- 
pared by the authors is labelled as showing a “dis- 
cernable pattern with characteristic variability”. The 
discernable patterns are lines overlaid on the plotted 
points without any statistical support. This would not 
be acceptable in an undergraduate ecology class, let 
alone a scholarly thesis. 

It is unfortunate that the authors make no reference 
to Lee et al. (1998), which has become the standard 
forest classification system for Ontario. It may be in- 
teresting to contrast the ecological land classification 
(ELC) of Lee et al. with Martin and Martin’s forest 
communities. If their data could be used to refine or 
correct the ELC system it would be far more useful 
than it is as a stand-alone study. 

The most interesting part of this book is the rela- 
tionship between forest types and their fauna. The 
authors have collated a great deal of their own data as 
well as data from published and unpublished sources. 
A more thoughtful analysis of this aspect of their 
study might produce a more enduring contribution to 
the study of forest ecology in Ontario. 


Literature Cited 

Lee, H., W. Bakowsky, J. Riley, J. Bowles, M. Puddister, P. Uhlig, 
and S. McMurray. 1998. Ecological land classification for 
Southern Ontario: first approximation and its application. Ontario 
Ministry of Natural Resources, Southcentral Science Section, 
Science Development and Transfer Branch. SCSS Field Guide 
FG-02. 


TYLER SMITH 


5900 rue Monkland, Apartment 10, Montreal, Quebec, 
H4A 1G1 Canada 


Les champignons des arbres de l’est de l’ Amerique du Nord 


By Bruno Boulet. 2003. Les Publications du Québec, Sainte- 
Foy, Quebec, Canada. 727 pages. $49.95 


The author and a number of colleagues have invest- 
ed a considerable effort to produce this detailed treatise 
on the principal wood-inhabiting fungi (mostly poly- 
pores) of eastern North America. The 19 x 24 cm, 
hardcover book is heavy but the binding is robust. The 
book contains nearly 500 color pictures, typically 6 or 
7 on a page, of very good quality. 

There are three principal sections. First there is a 
brief commentary on the vegetation of Quebec, espe- 
cially in relation to the geographic and host distribution 
of the wood-inhabiting fungi. Then 118 pages are 
devoted to a discussion of the state of our knowledge 
of the polypores, including their medicinal properties, 
edibility, and traditional and commercial uses. The role 
of these fungi as forest pathogens and their influence 
on the management of forests are reviewed and various 
tree defects caused by these fungi are depicted in color. 

The second section of the book introduces the reader 


to the taxonomy, nomenclature and classification of 
the polypores, as well as their distinctive macroscopic 
and microscopic features. The latter are in some in- 
stances essential in naming specimens. This section 
concludes with an explanation of the most common 
means of identifying and naming a specimen, i.e., the 
botanical dichotomous key, and a Key including over 
400 taxa. About 120 of the 400 are mushrooms that 
are briefly described in the key and 76 of them are 
shown in a color photo. Many are truly wood-inhab- 
iting fungi but a few are mycorrhizal (Lactarius and 
Russula) and others are incidental inhabitants of well- 
rotted woody debris. 

The third and the major part of the book presents 
essential information that allows recognition of 169 
species, observations on their biology and ecology, 
color pictures, and distribution maps. Readers should 
be cautious in their interpretation of the distribution 
maps because when I compared 8 maps with distri- 
butions in two references from the bibliography, 1.e., 


2004 


numbers 142 and 145, each map was deficient. Two 
specific examples are (1) reference 142, titled Alba- 
trellus in Michigan, includes A. caeruleoporus, A. 
confluens, A. cristatus and A. peckianus but Michigan 
is not shaded on the maps for those species, and (2) 
shading on the map for Sistotrema confluens covers 
southern Quebec and Nova Scotia but reference 145 
notes its presence in those provinces as well as in 
Michigan, North Carolina, New Hampshire, New York, 
Vermont and Wisconsin. 

Two new species, Auricularia americana Parm. et 
I. Parm. ex Audet, Boulet et Sirard and Polyporus long- 
iporus Audet, Boulet et Sirard, are proposed. Several 
species are reported for the first time in North America, 
for example, Antrodiella pallasii, Postia alni, P. folli- 
culocystidiata, P. ptychogaster, Phellinus cinereus and 
Polyporus tubaeformis. Although two names, Postia 
minisculoides and P. subpendula, are proposed as new 
combinations, if their basionyms, i.e., the initial name 


Book REVIEWS 


147 


given to the fungus and its place of publication, are 
not in the book the new combinations are not validly 
published. There is a picture (plate 15 D) labeled Punc- 
tularia strigosozonata, that shows a typical fruit body 
of the orange crust fungus, Phlebia radiata. 

The book concludes with a glossary containing ap- 
proximately 250 terms with their English equivalent 
and a definition that often includes a reference to a 
page where the term is used or illustrated, an index of 
French and English common names, an index of sci- 
entific names, and a bibliography of 450 entries. 

The book is recommended to mycologists, forest 
pathologists, forest ecologists, and naturalists. It is a 
significant contribution to our knowledge of the wood- 
inhabiting fungi of eastern North America. 


J. GINNS 


1970 Sutherland Road, Penticton, British Columbia V2A 8T8 
Canada 


Manual of Vascular Plants of Northeastern United States and Adjacent Canada: Second Edition 


By Henry A. Gleason and Arthur Cronquist. 2004. The New 
York Botanical Garden Press, 200th Street and Kazimiroff 
Boulevard, Bronx, New York 10458-5126 USA. 993 pages. 
U.S.$69.00. Cloth. 


The second edition of this most useful flora was 
first printed in 1991. Subsequent printings have taken 
place in 1993, 1996, 1998, 2000 and 2002. The seventh 
printing which has a slightly larger page (15 cm x 23 cm, 
rather than 14 cm x 21.5 cm) has a slightly larger type- 
face. The text pages have exactly the same page numbers 
as the earlier printings. Some corrections were made in 
the 1993 volume. In 1999 nineteen individuals con- 
tributed numerous corrections but these corrections 
could not be included in the 2000 and 2002 printings 
without having an electronic version. One was finally 
made for the 2004 printing. 

This new volume has an interesting but almost hid- 
den drawing of a Tulip-tree, Liriodendron tulipifera, 
on the front cover. The introductory pages start with a 
Table of Contents which includes a list of the families 


in taxonomic order with their page numbers. This is 
followed by a Foreword by Patricia K. Holmgren and 
Noel H. Holmgren and short bibliographies of Henry 
Allan Gleason and Arthur Cronquist by Noel H. 
Holmgren together with photographs of them. The 
Glossary which preceded the synoptic keys in the 
earlier printings now follows the main text. This is 
followed by the Index to Common Names and the 
Index to Scientific Names which in the earlier volumes 
were combined. 

This new volume is a most welcome step ahead 
with the numerous changes and corrections, and The 
New York Botanical Garden Press is to be congratulated 
even though it is still called the Second Edition. 


WILLIAM J. Copy 


Biodiversity, National Program on Environmental Health, 
Agriculture and Agri-Food Canada, Research Branch, Wm. 
Saunders Building, Central Experimental Farm, Ottawa, 
Ontario K1A 0C6 Canada 


The Wild Orchids of North America, North of Mexico 


By P. Martin. 2003. University Press of Florida, 15 Northwest 
15th Street, Gainesville, Florida. USA. $27.95 paper, 
U.S.$45.95 cloth. 


This book is a special version of a check list. The 
orchids included are all the recorded species found 
above the U.S.—Mexican border north to the Arctic 
and Greenland. This covers 223 species plus 24 sub- 
species and varieties. Additionally, this takes in 103 
growth and color forms, 24 hybrids, and introduced 
species. The species are arranged alphabetically by sci- 
entific name, so the first entry is the charming little 
Spotted Orchid Amerorchis rotundifolia. The author 
gives the genus, synonyms, misapplied names, typical 


common name, references and range. The author also 
adds any appropriate comments. For each species there 
is a5 x 7 cm photograph of the flower and a line draw- 
ing, generally of the whole plant. In some cases the line 
drawing is of the flower only which is a duplication 
that does not add information. 

In addition to the over 60 species that can be found 
in Canada, and the familiar genera (Platanthera, 
Cyprepedium etc.), there are several genera that | 
normally associate with the tropics (Vanilla, Laelia, 
Epidendrum). While many of these are escapees from 
cultivation, there are a number that are native species. 
Not surprisingly, most of these tropical epiphytes are 
to be found in Florida. Indeed, I was surprised to see 


148 


how many species of Vanilla (of vanilla ice-cream fame 
from Vanilla planifolia) that are native to Florida. 

The author uses the term “waif” to denote random 
individual occurrence. An example of a waif is Laelia 
rubescens, first seen in Florida in 1999. This abundant 
and attractive Central American species is a popular 
garden plant and a likely escapee. The accompanying 
photograph shows a white blossom. All the wild L. 
rubescens | have seen were pale lavender. 

The species coverage and their current status are 
both accurate and up-to-date. This book contains the 
split between the Spotted Orchid, Cyprepedium gutta- 
tum (Alaska and NWT), and the Yellow Spotted Orchid, 
C. yatabeanum (Kodiak Island). It includes the New- 
foundland orchis (Platanthera albida) of Newfoundland 
and Greenland. The book does not contain any habitat 
information, nor anything of the plants’ biology. Gen- 
erally, nothing is given on abundance and the distribu- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


tions are very generic (e.g., Manitoba east to New- 
foundland south to Texas and Georgia). It would be 
exciting to see an expanded version of this book con- 
taining descriptive text on habitat and biology, accom- 
panied by useable range maps. This would create an 
encyclopedia of North American orchids. In the mean- 
time, this book is a very handy reference and will make 
a good field guide. 

The book’s size precludes it fitting a pocket, but it 
will slip easily into your back pack. As well as the 
usual glossary and bibliography, the author includes 
a well-organized key. This, combined with the clear 
format, makes it a no-nonsense, practical guide for 
botanists, naturalists and orchid enthusiasts. 


Roy JOHN 


2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario 
K1iJ 6K5 Canada 


Trees of the Carolinian Forest: A Guide to Species, Their Ecology and Uses 


By Gerry Waldron. Boston Mills Press, 132 Main Street, 
Erin, Ontario NOB 1TO Canada. 274 pages. $24.95. 


The Carolinian zone of southern Ontario is home to 
more than 1600 plant species. It is also the most dense- 
ly populated area of the country. As a consequence, 
Canada’s most biologically diverse forests are also 
among the habitats most threatened by development. 
Anyone with an interest in the appreciation and con- 
servation of this natural heritage will do well to read 
Gerry Waldron’s Trees of the Carolinian forest. Mr. 
Waldron has succeeded in producing a beautiful book 
that is at once a pleasure to read and quite informa- 
tive. The subtitle, “A guide to species, their ecology 
and uses” is somewhat misleading — this is much more 
than a field guide. Waldron draws on a variety of his- 
torical and scientific sources to set the scene: what is 
the Carolinian zone? how is it related to other eco- 
logical regions? and how have successive human cul- 
tures altered this region? With the ecological context 
established, Waldron treats each of 73 tree species 
not as individuals but as members of a community. 

The book starts with an ecological history of the 
Carolinian zone, from glaciation through settlement 
by indigenous and European humans up to the present. 
While far from an presenting an exhaustive review of 
post-glacial colonisation, Waldron includes enough 
detail to distinguish his treatment from the usual cli- 
chéd summary that appears in books of similar scope. 
We learn of the massive hemlock dieback 5000 years 
back, and that beech was averaging 20 kilometres per 
century as it crept north. This is a fascinating subject, 
and it’s a shame that Waldron includes no references 
to his sources here. While in-text citations would be 
overly pedantic, including a few key references such 
as Pielou (1991) would be worthwhile. 

Waldron uses quotations from the journals of early 
settlers and survey crews to illustrate both the appear- 


ance of the “virgin” forest and the attitudes of Euro- 
peans to their new homeland. Once again, he piqued 
my curiosity, but in this case there are perhaps no read- 
ily accessible published sources he could refer the 
reader to for further information. 

A short discussion of the definition of the term 
“Carolinian” follows. Waldron covers the topic in five 
pages — a clearly presented summary of an important 
concept. This is typical of the book as a whole. The 
author quite capably distills complex ideas into simple 
language, without sacrificing accuracy in the process. 

With the context established, Waldron devotes some 
forty pages to a discussion of biodiversity, ecological 
communities, and our role in their protection, and 
especially, their restoration. He obviously brings a lot 
of experience to bear on the subject. Most books on 
restoration focus on technical details — how, where, 
and when to plant a tree, etc. Refreshingly, Waldron 
starts by examining why (and why not) to plant trees 
as part of a restoration. In a region where restoration 
of endangered prairie habitat often begins with the 
removal of trees planted as part of misguided natu- 
ralization programs, this is an important discussion. 
That said, he acknowledges the difficulty in pursuing 
a “do-nothing” approach, and offers suggestions for 
accelerating natural successional processes. These are 
presented as ideas to consider, not as ready-made pre- 
scriptions for restoration success. 

And so it is that on page 112 of this tree guide that 
the actual species treatments begin. Each species is 
allotted two pages. This includes the usual description 
of their habitat and appearance, with insights into their 
use in restoration and urban plantings. Unfortunately, 
Waldron has chosen to arrange the species alphabet- 
ically by common name. This may simplify things for 
the beginner, but it has the unfortunate consequence 
of separating walnut (Juglans nigra) from butternut 
(Juglans cinerea) and poplar from aspen (both Popu- 


2004 


lus). Taxonomic arrangements are a very useful tool 
for learning about the relationship between species, 
but only if we use them! 

The book is rounded out with 32 colour plates. 
These include a variety of habitat, habit, and close-up 
photographs of Carolinian trees. Some of them are 
quite instructive, such as the close-ups of the bark 
cross-sections of red and white elm, or the collection 
of acorns from different oak species. Others are simply 
beautiful images of the authors favourite organisms. 
One small improvement would be the inclusion of a 
scale for the close up shots. 

Taken as a whole, this book is an incredible resource 
for naturalists in southern Ontario. It will serve equally 


ENVIRONMENT 


Book REVIEWS 


149 


well as an introduction to the ecology of the Carolin- 
ian forest for the general naturalist and as an idea 
book for restorationists and land managers. I look 
forward to the publication of a companion volume on 
the shrubs of the Carolinian forest, hinted at in the 
introduction. 


Literature Cited 
Pielou, E. C. 1991. After the ice age: the return of life to glaciated 
North America. University of Chicago, Chicago. 


TYLER SMITH 


5900 rue Monkland, Apartment 10, Montreal, Quebec, 
H4A 1G1 Canada 


Good News for a Change: How Everyday People are Helping the Planet 


David Suzuki and Holly Dressel. 2003. Greystone Books, 
#201 — 2323 Quebec Street, Vancouver, British Columbia 
V5T 4S7 Canada. 399 pages. $24.95, U.S.$16.95 


This is a paperback re-issue of Suzuki and Dressel’s 
2002 book originally titled Good News for a change: 
hope for a troubled planet. Either way, the book is 
cleverly titled to attract readers who are tired of doom 
and gloom environmental books. The concept is praise- 
worthy: bring together in one place success stories of 
the many and varied ways people around the world 
are changing the way they do things, for the benefit 
of their communities, their environment and their 
offspring. At the same time, the authors do not shy 
away from the bad news. Yes there are wonderful 
changes taking place but they are up against very seri- 
ous problems and a frightening inertia embedded in 
our social and economic systems. 

The authors have set a tremendous challenge for 
themselves in tackling the full sweep of environmental 
problems we face and in trying to represent emerging 
solutions from around the planet. They organize this 
unwieldy subject thematically with chapters on busi- 
ness practices, democracy, biodiversity, water, food, 
forests, fisheries, and air pollution. The final chapter 
“Breaking out of the Box” addresses the effect of our 
global culture on humans as natural beings and the 
importance of reconnecting what we do with our deep- 
est values. Most chapters focus on one or two case 
studies examined in some depth with additional exam- 
ples of related projects in other parts of the world. 
Examples are drawn from village co-operatives, family- 
run businesses, multi-national corporations, farms, in- 
digenous societies and non-profit groups from such 
diverse places as India, Africa, Germany, Brazil, USA, 
Indonesia and many others. Also included are a list 
of organizations to contact, detailed endnotes, a brief 
bibliography and a reasonably detailed index. The 
book is a little lax in explaining the political back- 
ground to some of its examples, especially Canadian 


ones, and a map plotting the locations for major exam- 
ples would have been beneficial. 

Throughout Good News there is an admirable effort 
to synthesize, and several key themes are followed. 
Aligning our activities with natural environmental 
systems results in double dividends — the costs are 
lower, the benefits are greater and extend beyond eco- 
nomics. A resource is used sustainably when it is man- 
aged by a stable community that is economically de- 
pendent on it and exerts local control and local 
ownership. Sustainable practices must be tailored to 
the specific situation; the people who follow them are 
humble and are constantly learning from their mis- 
takes. Many small projects are more effective and more 
efficient than a single large one. We are all indigenous 
to this planet. 

If I have any quibble with this book it is that there 
isn’t enough of it. I want to know more about how 
Judy Wicks manages to make a living (and a rather 
substantial one) running a social activist restaurant. I 
want to understand how a small check dam on an in- 
termittent stream in India can reverse desertification 
caused by forest removal. I want to hear from the 
efforts that haven’t worked, because I don’t believe 
as easy at the book makes it seem. There is no question 
we need more books like this, perhaps ones targeted 
at specific issues. Henry Mintzberg (1994) argues for 
the importance of identifying “emergent strategies” 
(new strategies that emerge spontaneously at any level 
of an organization) and helping them spread. That in 
essence is what Suzuki and Dressel have done. 


Literature Cited: 
Mintzberg, H. 1994. The Rise and Fall of Strategic Planning. Free 
Press, Simon and Schuster, New York, USA. 454 pages. 


CAROLYN SEBURN 


Seburn Ecological Services, 920 Mussell Road, RR 1, Oxford 
Mills, Ontario KOG 1S0 Canada 


150 


Whose Bird? 


By Bo Boelens and M. Watkins. 2004. Yale University Press, 
P.O. Box 209040, New Haven, Connecticutt 06520-9040 
USA. x + 400 pages U.S.$35 paper. 


Have you ever been curious as to who was the 
Bonaparte of Bonaparte’s Gull fame? Was it really 
the feared Corsican tyrant? Now you can get a book 
that will answer that question and many more. The 
authors have researched about 1400 people who have, 
at one time or another, given their names to bird spe- 
cies. There is a cameo biography for each individual. 
As the book gives the accepted English names, there 
is a very slight bias to the coverage. European and 
American naturalists dominate the list, but to be fair 
Professor Ijima (jima’s Warbler, Phylloscopus tjimae) 
gets his credit. Montezuma does not fare as well. He 
gets a single sentence biography stating he was Em- 
peror of the Aztecs. Actually, Montezuma I, Emperor 
of Mexico, was a remarkable man. He was a victorious 
general before he succeeded his uncle as emperor in 
1436. He is only recognized for his Quail, with no 
mention of his strikingly handsome Oropendola. In 
contrast, Californian Alberto Treganza (Treganza’s — 
now Great Blue — Heron) gets almost a whole page. 

Many of the names have fallen by the wayside. The 
Bancroft’s Night Heron is now called by the more 
descriptive name of Yellow-crowned Night Heron. The 
loss of some names is. a benefit. | would have trouble 
spelling and pronouncing Sjéstedt (Sjostedt’s Owlet, 
now Barred Owlet). No disrespect to Mr. Helmut Sick 
(Sick’s Manakin), but I think the Golden-crowned 
Manikin would prefer its new name. However, I am 
sad to see Lear’s Macaw is now an Indigo Macaw. 
Edward Lear, mostly known for his nonsense poems, 
was a remarkable Victorian bird artist who deserves a 
memorial. William MacGillivray’s name lives on as 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


MacGillivray’s Warbler (Oporornis tolmia) but his 
son’s bird, MacGillivray’s Petrel, is now the Fiji Petrel, 
but scientifically is Pseudobulwaria macgillivrayi (The 
generic name honours the Reverend James Bulwer.). 

Some of these names are still current. Thekla Lark, 
named after the dying daughter of a German natural- 
ist is still a Thekla Lark. Of the 17 species named for 
the pioneering Dutch ornithologist Coenraad Temmink, 
six still remain. I am delighted that the Adélie Penguin’s 
name remains intact as it really suits this charming 
bird. Be thankful that Admiral Jules-Sebastien-César 
Dumont D’Urville named it for his wife and not 
himself. 

There are some odd stories in the book. I was par- 
ticularly titivated by the tale of Colonel George 
Montagu (Montagu’s Harrier) who was cashiered from 
the British army for “provocative marital skirmishing” 
and thereby took up the study of science. I will never 
look at the harrier the same way again! 

There are some names that are conspicuously absent. 
Roger Tory Peterson, Peter Scott (Scott’s Oriole is 
named for Winfield Scott) and James Fisher were 
probably born too late in history. Linneus or Linné, 
who invented the nomenclature system, was alive in 
the right era, but remains un-honoured. 

As well as the cameos, the authors have collected 
together those people who shared a characteristic in 
common, such as diplomats, Germans or those who 
died of gunshot. Many of the entries have a black- 
and-white portrait of the person featured. For those of 
you who love trivia or are keen naturalists preferably 
both, this is a fun book. 

Roy JOHN 


2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario 
K1J 6K5 Canada 


Forest Dynamics and Disturbance Regimes: Studies from Temperate Evergreen-Deciduous 


Forests 


By Lee E. Frelich. 2002. Cambridge University Press, 40 
West 20th Street, New York, New York 10011-4221 USA. 
266 pages. GBP50 (U.S.$80). 


Disturbance is ubiquitous in forest ecosystems. 
Forested landscapes are best viewed as an integration 
of climatic, biotic, edaphic and geomorphic processes 
that determine the character of disturbance events 
occurring Over a wide range of temporal and spatial 
scales. Disturbed by the extremes of either catastroph- 
ic, stand-replacing events that may include fire, in- 
sect outbreak, and extensive windthrow, or periodic, 
small-scale gap processes mediated by fungal patho- 
gens, forests are in constant flux when viewed from a 
landscape perspective. Such a wide range in the per- 
iodicity, intensity and scale of disturbance events, 
and the diversity of bio-edaphic interactions create a 
complex, fluid, heterogeneous landscape. 


Lee Frelich, founder and director of the University 
of Minnesota Center for Hardwood Ecology, intro- 
duces the reader to the significant disturbances that 
have shaped, and continue to shape, the hemlock- 
hardwood forests of the northern regions of the Lake 
States (Minnesota, Wisconsin, and Michigan). For 
the past two decades Dr. Frelich has dedicated himself 
to understanding the stand- and forest-level dynamics 
of these deciduous-to-boreal transition forests. He 
forms part of a long tradition of university and govern- 
ment (United States Forest Service) forest ecology 
research, much of which is scattered in scientific jour- 
nals and government reports. Forest Dynamics and 
Disturbance Regimes provides for the first time, in an 
engaging, well-illustrated, and synthetic format, the 
fruit of this rich research legacy. 


2004 


“Under what conditions do forests change or stay 
the same?” Thus might one summarize the intent of 
this book. Introductory chapters set the scene by des- 
cribing the Great Lakes temperate forests and their 
disturbance regimes dominated by fire, wind, insect 
outbreaks and mammalian herbivory. Of significance 
to the practicing forest ecologist will be the chapter on 
sampling and interpretative techniques used to detect 
and interpret forest disturbance regimes. Emphasis is 
placed on the use of tree radial increment patterns as 
a valuable source of insight into stand disturbance 
history. 

Frelich emphasizes the critical role played by dis- 
turbance in both stand development and forest suc- 
cession. He properly distinguishes between stand 
development and succession, both of which are often 
confused in the literature. Disturbance will always ini- 
tiate a new cycle of stand development in the regen- 
erating, post-disturbance forest. However, disturbance 
may or may not initiate a species change or a new 
successional sequence. 

Consideration is also given to the differing effects 
of disturbance at both the stand- and landscape-level. 
This distinction is important, especially given the wide 
temporal and spatial scales at which disturbances may 
occur. Furthermore, instability at the stand level may 
be interpreted as stability at the landscape level. Inter- 
pretation often depends on the scale of investigation. 

A particular strength of this work is Frelich’s ability 
to engage the complex interaction of different distur- 
bances. Frelich not only introduces the wide diversity of 
temporal and spatial patterns of forest change, but even 
more importantly, highlights often counter-intuitive 
insights into forest change and continuity. I found the 
following particularly noteworthy: (1) the nonlinear 
response of forest species composition to disturbance 
severity, (2) the cause and development of patchy hard- 
wood-softwood mosaics, (3) clarifying taxonomy of the 
concept of old-growth, (4) how different forest types 


BooK REVIEWS 15] 


can exist on relatively homogeneous sites, and (5) the 
multiple successional pathways open to any particu- 
lar forest type. Frelich’s final chapter summarizes the 
notion of forest stability. It provides conceptual models 
of forest response to disturbance, 3-D models of suc- 
cession in different forest types, and a final classifica- 
tion of four different types of forest landscape. 

This work is particularly important as humans con- 
tinue to “disturb” forests, especially by commercial 
forestry. Before any claims can be made about the 
desirability of the changes created by human interven- 
tions, it is essential to properly comprehend the range 
of natural forest disturbance regimes and the associ- 
ated changes in forest structure and tree species com- 
position. 

The book addressees the scientific community and 
would properly of greatest interest to forest ecologists 
and all students of forest change. The judicious mix 
of empirical case studies, hypothetical examples and 
conceptual models helps the reader to think “beyond 
the box.” The many line drawings, flow charts and 
black-and-white photographs help to clarify the dif- 
ferent concepts. 

As one is reminded in the subtitle, this book focuses 
exclusively on the temperate evergreen-deciduous for- 
ests (of the Lake States). While it is certain that many 
of the concepts developed from research in this forest 
type are applicable to other forest types, it is wise to 
resist any quick and easy transfer of ideas. Forests grow 
in conditions that span a wide ecological spectrum, a 
situation that often resists our human tendency to 
categorize and classify. Be that as it may, this book 
provides rich and substantive insight into this well- 
studied — and much-loved — forest region at the decid- 
uous-boreal interface. 


JOHN McCartny, S.J. 


St. Mark’s College, University of British Columbia, 5935 
Iona Drive, Vancouver, British Columbia V6T 1J7 Canada 


Snowshoes & Spotted Dick; Letters from a Wilderness Dweller 


By Chris Czajkowski. 2003. Harbour Publishing Co. Ltd., 
PO Box 219, Madeira Park, British Columbia VON 2HO 
Canada. 298 pages. Paper U.S.$24.95 


This book is a one-sided account of the wilderness 
experiences of a woman of extraordinary independ- 
ence, as she writes letters to a friend named Nick. 
Chris Czajkowski is an author and wilderness guide 
who has chosen a life of relative and geographic iso- 
lation on remote property where she lives three quarters 
of the year in the coast mountains of British Colum- 
bia, 480 kilometres north of Vancouver. There is no 
telephone or mail delivery and the radio only works 
on clear days. The computer used while writing the 
book draws the electricity required to function from 
solar panels. The visitors to the eco-tourism business 
“Nuk Tessli Alpine Experience” are brought in by float 


plane, along with any supplies and news from the 
outside world. 

This, the fourth book by the same author, is her 
account of the incredible effort to build a third cabin on 
the property. Its format of letters scribed to a friend in 
Germany is generously interspersed with some textual 
description, a few black-and-white pictures and hand- 
drawn sketches. 

The reader is drawn in by expressive language evok- 
ing in the imagination pictures of the breathtaking 
vistas, and natural surroundings so that you feel almost 
as though you have been there, perhaps as a visitor to 
“Nuk Tessli”. The added mystery of “what is Spotted 
Dick?” is a cute grab but is just a little overdone. 

The easy flow of the language makes the 298 pages 
an easy read, despite the occasional construction jargon. 


sy 


The actual process of building a log cabin in the woods, 
particularly in such a remote location, is fascinating. 
The cabin is built in several steps, over numerous 
months, using hand tools, chainsaws, and, occasion- 
ally, some friends. 

There are a few drawbacks from the enjoyment of 
this book though, especially without having read the 
first three. The first is that in this book there are a 
number of interesting stories that are alluded to but 
not told in detail because they’ve been recorded in 
the previous books. The second is that the letters are 
all to Nick and because it does not include the letters 
written from him, it feels a bit like being in the same 
room with someone on the phone — you hear half the 
conversation and although you get the gist of the 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


whole, you feel the missing of the other half. The third 
and most perplexing is the lack of description of the 
characters or their relationship with each other. The 
reason this is perplexing is because the author herself 
describes this criticism received from a book review- 
er — and then simply passes it off with “The trouble 
is, I live with the people I write about. Experience 
has shown that no matter how innocuous a portrait I 
paint, the subject will find something about which he 
or she is unhappy.” While I am sure this could be 
true, the book would earn a wider audience if it were 
not written for those people alone. 

DAWN BURNETT 


Jacques Whitford Limited, 1 Union Street, Elmira, Ontario 
N3B 3J9 


Natural Grace: The Charm, Wonder, & Lessons of Pacific Northwest Animals & Plants 


By William Dietrich. 2003. University of Washington Press, 
1326 Fifth Avenue, Suite 555, Seattle, Washington 98101- 
2604 USA. 236 pages. U.S.$19.95 


Natural Grace is a contemporary example of the 
medieval book of beasts called a bestiary. Bestiaries 
were very popular during the Medieval Period, and 
focused on the life of nature as a model or paradigm 
for human behavior. These books were not scientific 
in the contemporary sense of the term, but instead 
combined religious and moral teaching with a close 
observation of nature, zoological commentaries, and 
fabulous and fictitious creatures. Rather than being 
studied in and of itself, nature was considered symbol- 
ic of both the virtue and moral life of human beings. 
While Deitrich’s work certainly provides the reader 
with a more sophisticated scientific understanding of 
nature, the subtitle of the book betrays its “bestiary” 
lineage as the reader is invited to appreciate the charm, 
wonder, and lessons of a variety of creatures and 
natural phenomena. 

This collection of essays is adapted from William 
Dietrich’s popular articles in the Seattle Times’ North- 
west Magazine. The author divides his work among 
four themes: (1) the common and ubiquitous creatures 
that we often take for granted, (2) the itty-bitty world 
that ranges from soil to spiders, (3) the rhythmic, 
clockwork world of tectonics and tides, and finally, 
(4) the iconic symbols of the Pacific Northwest includ- 
ing, among others, the bald eagle and the killer whale 
or orca. 

Dietrich is gifted with the contemplative eye. Follow 
his gaze and you will enter a world of delight and 
amazement. We live side by side with other creatures 
and are immersed within the workings of nature. But 
often, we are oblivious to the ways of our environs, so 
intent and fascinated are we with our own creations 
and fabrications. Dietrich pries apart our fabricated 
world and invites us to direct our gaze out beyond the 
confines of culture. He calls us to a long loving gaze 
on the other of nature. 


This contemplative gaze is a virtue that strengthens 
with time and practice. Dietrich is a patient and humor- 
ous guide, gently leading the reader to grow in knowl- 
edge of the natural world; knowledge, not simply of 
the biological or physical facts, but of the beauty, sig- 
nificance, and yes, even mystery inherent in the sub- 
ject of our gaze. Of special interest is the manner by 
which the author weaves together culture and nature. 
Whether it be the forces of nature that define the 
boundaries of human culture and provide the myriad 
free ecological services that allow us our cities and 
farms, or the manner in which particular species have 
become embedded in our folk and cultural lore, the 
reader is left with the knowledge that we are depend- 
ent on the “others” of nature far beyond our physical 
survival. 

Human ignorance instills fear, and fear breeds vio- 
lence and destruction. Natural Grace sets a contrary 
path. Knowledge of the other, of nature, can lead to 
intimacy, and intimacy can be the foundation of love. 
Love, in turn, leads to mutuality, care, and concern for 
the other. Dietrich’s melodious writing can indeed lead 
one to greater love of nature and hopefully, action on 
behalf of nature. 

This book is a lovely blend of nature and culture, 
of scientific vulgarization and social lore. You will 
learn much about the natural history of the Pacific 
Northwest. As well, I think, you will learn about how 
we (and maybe you yourself) relate to the natural 
world, and how that relationship has changed with 
time, and how it may or must change in the future. 

Given Dietrich’s place of habitation (Seattle), this 
book focuses on the State of Washington, with excur- 
sions south to Oregon and north to British Columbia. 
So, if you call the Pacific Northwest your home, you 
will learn much about your “neighbours.” However, 
regardless of one’s geography, Dietrich’s work is more 
a work of perspective, rather than of content. His 
actors live and work in the Pacific Northwest, but the 
perspective shared is universal. No matter where you 


2004 


live, the long, loving, contemplative gaze on the real 
is always a possibility. 

The 12" century Cistercian monk, Saint Bernard 
of Clairvaux, once said; “I have discovered that you 
will find far more in the forests than in books; trees 
and stones will teach you that which you cannot learn 
from any master.” Turn a discerning eye to the crea- 


Book REVIEWS 


153 


tures and processes that surround you and you will 
enter a world of marvel and beauty. Let Natural Grace 
be your companion and guide. 


JOHN McCarthy, S.J. 


St. Mark’s College, University of British Columbia, 5935 
Iona Drive, Vancouver, British Columbia V6T 1J7 Canada 


Tales from the Underground: A Natural History of Subterranean Life 


By David W. Wolfe. 2001. Perseus Publishing, 5500 Central 
Avenue, Boulder, Colorado 80301 USA. 221 pages. $27.50 


Tales from the Underground is a MUST read for 
any naturalist interested in what’s happening under 
our feet. The book is engaging, well-written, and filled 
with information that is truly awesome. 

In the introduction, author David Wolfe, Associate 
Professor of Plant Ecology at Cornell University, in- 
vites his “subterranean-impaired” readers to use their 
imaginations and join him on a dive trip into a mys- 
terious world scientists are only just beginning to un- 
derstand. Wolfe launches the journey with a description 
of the various soil “profiles”, also known as layers or 
horizons, and the organisms that inhabit them. Some 
of the life forms I had never heard of; others I had 
never thought about in the way Wolfe describes them. 

Take plants, for example. Wolfe starts by praising 
their display of foliage aboveground. He then points 
out that plants are unique because they simultane- 
ously inhabit both the surface and sub-surface realms. 
He calls them the “great mediators between the two 
realms”, and explains their functions both above and 
below the ground, with a focus on their buried other 
half: roots. 

In Part I of the book, “Ancient Life’, Wolfe covers 
a lot of ground. He discusses theories of the origin of 
life, the basic elements necessary for the evolution of 
life, Earth’s advantage as a life-generating planet, and 
speculation pointing to the subterranean realm as the 
place where life began. He also introduces “extremo- 
philes”, creatures that live in severe conditions thou- 
sands of feet underground, where pressure and tem- 
peratures are extraordinarily high, and where there is 
neither light nor oxygen — the postulated setting for 
the beginning of life. 

Wolfe also describes Carl Woese’s discovery of the 
amazing genetic diversity of these extremophiles, and 
discusses how that discovery changed the universal 
tree of life. Originally composed of bacteria, plants, 
fungi, animals and protozoa, the tree of life now has 


only three branches — bacteria, archaea (the extremo- 
philes) and eukarya (with plants and animals as two 
small twigs). It’s a fascinating and humbling picture. 

In Part I, “Life Support for Planet Earth”, Wolfe 
dedicates an entire chapter to Charles Darwin and the 
lasting influence of the renowned scientist’s meticu- 
lous and painstaking work. Wolfe also portrays certain 
inconspicuous subterranean creatures and the impor- 
tant roles they play in cycling essential elements and 
facilitating the flow of energy — processes that sustain 
life on this planet. He also discusses the dual roles 
played by soil with respect to plant and animal dis- 
eases; some soil microbes cause diseases, while others 
provide powerful antidotes. 

The third part of the book, “The Human Factor”, 
describes the enormous footprint of Homo sapiens on 
the planet. Wolfe tells the sobering, appalling story 
of human activities and their devastating impacts on 
wildlife such as prairie dogs, black-footed ferrets, and 
burrowing owls, and on soil integrity in general. Des- 
criptions of soil erosion and the 1930s dust bowl crisis 
are particularly gripping. 

Yet despite the doom and gloom, Wolfe ends on an 
optimistic note. The very last sentence of the book 
reads, “It is my hope that as more of us become aware 
of the life beneath our feet, and its relevance to our 
well-being, we will be inclined to work together to 
maintain the biological integrity of the underground, 
and preserve some of what we find there for future 
generations” — words we have heard before in one form 
or another, but good to see repeated in this context. 

Tales from the Underground is David Wolfe’s first 
book, and I hope it isn’t his last. He has a pleasant and 
easy-to-read writing style, and manages to convey sci- 
entific concepts so that even a non-scientific type like 
me can understand. Tales from the Underground has 
found a permanent place on my bookshelves. 


R. SANDER-REGIER 


RR5 Shawville, Quebec JOX 2YO Canada 


154 


NEw TITLES 


+Available for review * Assigned for review 


Zoology 
* Amphibian Decline: An integrated analysis of multi 
stressor effects. By Greg Linder. 2004. SETAC North Amer- 
ica, 1010 North 12th Avenue, Pensacola, Florida. U.S.$98 
Cloth. 


Annotated Checklist of Birds of Chile. By M. Marin. 2004. 
Lynx Edicions, Barcelona, Spain. 144 pages. U.S.$15.75 [in 
English and Spanish]. 


Biology of Sharks and their Relatives. By J. Carrier, J. Mu- 
sick and M. Heithus. 2004. CRC Press, 2000 NW Corporate 
Boulevard, Boca Raton, Florida. U.S.$99.95. 


* Birds of Belize. By H. L. Jones. 2004. University of Texas 
Press, Austin, Texas. 317 pages, U.S.$60.00 Cloth, $34.95 
Paper. 


Birds of South Asia. By P. Rasmussen and J. Anderton. 2004. 
Lynx Edicions, Barcelona, Spain. U.S.$95 Cloth. 


Birds of Spain. 2004. Lynx Edicions, Barcelona, Spain. 296 
pages. U.S.$49 [in English and Spanish]. 


Birds of the Raincoast. By H. Thommasen and K. Hutchings. 
2004. Harbour Publishing, P.O. Box 219, Madeira Park, Brit- 
ish Columbia. 260 pages. U.S.$24.95. 


The Exotic Amphibians and Reptiles of Florida. By W. 
Meshaka, B. Buttergield and J. Hauge. 2004. Krieger Pub- 
lishing P.O. Box 9542 Melbourne, Florida 32902-9452. 166 
pages. U.S.$34.50. 


* Hawks and Owls of the Great Lakes Region & Eastern 
North America. By C. Earley. 2004. Firefly Books Ltd., 3680 
Victoria Park Avenue, Toronto, Ontario, M2H 3K1. 128 pages. 
$16.95 Paper, $24.95 Cloth. 


+ Orca — Visions of the Killer Whale. By Knudtson. 2004. 
Greystone Books, Vancouver, British Columbia. xvii +110 
pages, $19.95. 


Pacific Reef and Shore. By R. Harbo. 2004. Harbour Pub- 
lishing, P.O. Box 219, Madeira Park, British Columbia. 80 
pages. U.S.$9.95. 


Reproductive Biology and Early Life History of Fishes in 
the Ohio River Drainage. By T. Simon and R. Wallus. 2004. 
CRC Press, 2000 NW Corporate Boulevard, Boca Raton, 
Florida U.S.$119.95. 


* Self-Portrait with Turtles: A Memoir. Edited by David M. 
Carroll. 2004. Houghton Mufflin Co., 222 Berkeley Street, 
Boston, Massachusetts 02116. 192 pages. 40 black-and-white 
line drawings and halftones. $23.00 Hardcover. 


Botany 
* Arboretum America: a Philosophy of the Forest. By Diana 
Beresford-Kroeger. 2003. University of Michigan Press, 
839 Greene Street, P.O. Box 1104, Ann Arbour, Michigan. 
196 pages. U.S.$52.50. 


Botanical Latin: Fourth Edition. By William T. Stearn. 
1992 [Re-issued as a paperback 2004] Timber Press, 133 SW 
2nd Avenue, Suite 450, Portland, Oregon, USA. 560 pages. 
Paperback, U.S.$29.95. 


* Manual of Vascular Plants of Northeastern United States 
and Adjacent Canada. By H. Gleason. 2004. New York 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Botanical Gardens, 200" Street and Kazimiroff Boulevard, 
Bronx, New York. 993 pages. U.S.$69.00. 


The Orchid in Lore and Legend. By Luigi Berliocchi. 2000 
[Re-issued as a paperback 2004] Edited by Mark Griffiths. 
2000 Timber Press, 133 SW 2nd Avenue, Suite 450, Portland, 
Oregon, USA. 200 pages. Paperback, U.S.$19.95. 


The Secrets of Wildflowers. By J. Saunders. 2004. The Lyons 
Press, 246 Goose Lane, P.O. Box 480, Guilford, Connecticut. 
304 pages. U.S.$24.95. 


Wildflowers of the Seacoast in the Pacific Northwest. By 
J. Trelawny. 2004. Harbour Publishing, P. O. Box 219, 
Madeira Park, British Columbia. 80 pages. U.S.$12.95. 


Environment 
Arctic National Wildlife Refuge. By S. Banerjee. 2004. 
Harbour Publishing, P.O. Box 219, Madeira Park, British 
Columbia. 176 pages. U.S.$36.95. 


+ The Bird Almanac. By D. Bird. 2004. Key Porter Books, 
70 The Esplanade, Toronto, Ontario, Canada. xvii + 460 
pages, not illustrated. $24.95. 


* Dancing at the Dead Sea — Tracking the World’s Envi- 
ronmental Hotspots. By Alanna Mitchell. 2004. Key Porter 
Books, 70 The Esplanade, Toronto, Ontario MSE 1R2 Canada. 
259 pages. $26.95. 


Estuarine Research, Monitoring and Resource Protection. 
By M. Kennish. 2004. CRC Press, 2000 NW Corporate 
Boulevard, Boca Raton, Florida. U.S.$119.95. 


Important Transboundary Belarussian-Lithuanian and 
Lithuanian-Russian Wetlands. By L. Raudonikis, A. Skura- 
tovich, L. Baléiauskas, E. Drobelis, D. Grishanov. 2003. 
Botanikos institutas, Vilnius, Lithuania. 


* The History of Ornithology in Virginia. By D. Johnston. 
2004. University of Virginia Press, P.O. Box 400318, 
Charlottesville, Virginia. x + 219 pages. U.S.$35. 


The Lewis and Clark Columbia River Water Trail: A Guide 
for Paddlers, Hikers, and Other Explorers. By Keith G. 
Hay. 2004. Timber Press, 133 SW 2nd Avenue, Suite 450, 
Portland, Oregon, USA. 264 pages. Paperback, U.S.$19.95. 


* Politics of the Wild - Canada & Endangered Species. 
Edited by K. Beazley and R. Boardman. 2004. Oxford 
University Press, 70 Wynford Drive, Don Mills, Ontario, Can- 
ada. x + 254 pages, not illustrated. $29.95. 


* A Primer of Ecological Genetics. By J. Conner and D. 
Hartt. 2004. Sinauer Associates, Box 407, Sunderland, Massa- 
chusetts, USA. 207 pages. U.S.$34.95. 


+ Resource and Environmental Management in Canada. 
Edited by B. Mitchell. 2004. Oxford University Press, 70 
Wynford Drive, Don Mills, Ontario. x + 600 pages, $29.95. 


* Whose Bird? By B. Boleus and M. Watkins. 2004. Yale 
University Press. 384 pages. U.S.$35.00 paper. 


Children’s books 
Saving Birds: Heroes around the world. By P. Salmansohn 
and S. Kress. 2004. Tilbury House, Gardiner, Maine. 40 pages. 
U.S.$16.95. 


King Of Fish: The Thousand Year Run of Salmon. By E. 
Bauer. 2003. Voyageur, Stillwater, Minnesota. 160 pages. 
U.S.$29.95. 


News and Comment 


Point Pelee Natural History News 3(4) 


The Winter 2003 issue, volume 3, number 4, pages 53-68, 
contains: Point Pelee: Its Evolution and Structure (Alan S. 
Trenhaile) — Noteworthy Bird Records: September to Novem- 
ber 2003 (Alan Wormington) — More Cave Swallows: The 
Fall of 2003 (Alan Wormington) — Point Pelee Christmas Bird 
Count: December 15, 2003 (Sarah E. Rupert) — Fox Squirrels 
at Point Pelee: 1977, 1990 and 1997 (Alan Wormington) — 
In the Field (Ross’s Gull: Second Record for Point Pelee; 
“Harlan’s” Red-tailed Hawk: New to Point Pelee). 

The Point Pelee Natural History News was edited by Alan 
Wormington (e-mail: wormington@juno.com). Editorial Assis- 


COSEWIC Assessment Results November 2003 


This update on Canadian Species at Risk issued by the 
Committee on the Status of Endangered Wildlife in Canada 
(COSEWIC) is 44 pages and contains: About COSEWIC, 
(mandate, membership, definitions) — Summary Tables 
[COSEWIC species designated in five “risk” categories and 
in Not at Risk, and Data Deficient categories (Tables 1-3)] — 
Results of November 2003 COSEWIC meeting (Tables 4-5) 
— COSEWIC Assessment Results — Explanation of sym- 
bols — Species examined in five “risk” categories (Table 6), 
Not at Risk (Table 7), and Data Deficient (Table 8) — Record 


tants were Gordon D. Harvey and Michelle T. Nicholson; web 
site www.wincom.net/~fopp/Natural_History_News.htm. This 
is the last issue as the Board of Directors of the publisher, 
The Friends of Point Pelee, has decided to terminate the pub- 
lication due mainly to the lack of material being submitted. 
All past issues are available in the Nature Nook Book Store 
in Point Pelee National Park’s Visitor Centre or by mail from 
the Friends of Point Pelee, 1118 Point Pelee Drive, Leaming- 
ton, Ontario N8H 3V4 Canada; phone 519-326-6173; e- 
mail: fopp@ wincom.net. 


of Status Re-examinations — Record of Name Changes. 
Listed are 12 extinct, 21 extirpated, 160 endangered, 108 threat- 
ened, and 140 of special concern. Of the 441 forms in these 
categories, 65 are mammals, 58 birds, 31 reptiles, 19 amphib- 
ians, 78 fishes, 16 arthropods, 19 molluscs, 140 vascular plants, 
9 mosses, and 6 lichens. In addition, 32 forms have been 
considered and found not at risk, and 29 to be data deficient. 

This publication is available from COSEWIC Secretariat, 
Canadian Wildlife Service, Environment Canada, Ottawa, 
Ontario K1A 0H3. See Web site: http://www.cosewic.gc.ca. 


Froglog: Newsletter of the Declining Amphibian Populations Task Force (61) 


Number 60, February 2004. Contents: A Workshop on the 
Amphibians of Madagascar at Giand (Franco Andreone) — 
Amphibian Colonization of Mitigation Wetlands in Nebraska 
(Mark M. Peyton) — Annual Meeting Report from Canada 
(David Galbraith) — A Study of the Plethodontid Salamander 
Populations at Raccoon Creek State Park, Pennsylvania (Jen- 
nifer D. Haney and Mary S. Kostalos) — Reports on DAPTF 
Seed Grants — Froglog Shorts — New Book on Amphibian 
Declines (Amphibian Declines: An Integrated Analysis of Mul- 
tiple Stressor Effects. 2003. SETAC Press, Pensacola, Florida: 
http://www.setac.org/pubs. html). 


Froglog is the bi-monthly newsletter of the Declining 
Amphibian Populations Task Force of The World Conservation 
Union (IUCN)/Species Survival Commission (SSC) and is 
supported by The Open University, The World Congress of 
Herpetology, and Arizona State University. The newsletter is 
edited by John W. Wilkinson, Department of Biological Sci- 
ences, The Open University, Walton Hall, Milton Keynes, 
MK7 6AA, United Kingdom; e-mail: daptf@open.ac.uk. Fund- 
ing for Froglog is underwritten by the Detroit Zoological 
Institute, P. O. Box 39, Royal Oak, Michigan 48068-0039, 
USA. Publication of issue 61 was also supported by Peace 
Frogs www.peacefrogs.com and by RANA and the US 
National Science Foundation grant DEB-0130273. 


The Boreal Dip Net/L’Epuisette Boreal: Newsletter of the Canadian Amphibian and Reptile 
Conservation Network — Reseau Canadien de Conservation des Amphibiens et des Reptiles 


8(1) January 2004 


Contents: Editor’s Note: thanks to sponsors TD Bank 
Financial Group, TD Friends of the Environment, The Pelee 
Island Winery, Parks Canada [Kerrie Serben] — A note 
about our acronym [David Galbraith] — CARCNET/RECCAR 
2003: The Pelee Isand Experience [David Galbraith] — 
Summary of Species at Risk Proceedings from the 8th Annual 
Meeting of CARCNET/RECCAR and the 3rd Annual Pelee 
Island Winery Endangered Species Festival (ESF) [Kim 
Barrett and Ben Porchuk] — The Lake Erie Watersnake: A 
23-year Perspective on Microevolution, Population Status, 
and Recovery Planning [Richard B. King] — Loving Nature 


to Death [Ron Brooks] — Another Leap into Learning [Sara 
L. Ashpole] — Books to Get: Turtles [Anita Baskin-Salzberg 
and Allen Salzberg]; A guide to creating vernal ponds — 
Endangered Northern Leopard Frog Reintroduced into East 
Kootenay After 20 Year Disappearance — Frogs and Toads 
of Canada CD — Current CARCNET/RECCAR Board of 
Directors — New Species of Salamander Found in Revel- 
stoke [Larry. Halverson] — Paper on Traffic Mortality and 
Turtles [reviewed by David Seburn] — Coming soon... 
CARCNET 2004 September 24-27 Edmonton — Member- 
ship in CARCNET/ RECCAR. 


LDS) 


156 


For information on membership in the Canadian Amphibian 
and Reptile Conservation Society/Reseau Canadien de Con- 
servation des Amphibiens et des Reptiles ($10 students, $16 
non-students] contact Bruce Pauli, Canadian Wildlife Service, 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


National Wildlife Research Centre, Carleton University, Raven 
Road, Ottawa, Ontario K1A 0H3. Web site: http://www.carc 
net.ca/ 


Annotated List of the Arctic Marine Fishes of Canada 


In 2004 Fisheries and Oceans Canada, Central and Arctic 
Region, Winnipeg, Manitoba, printed a limited number of 
copies of the Annotated List of the Arctic Marine Fishes of 
Canada by B. W. Coad and J. D. Reist as Canadian Manu- 
script Report of Fisheries and Aquatic Sciences 2674, iv + 
112 pages. A primary list covers the 189 species represent- 
ing 115 genera in 48 families of marine and anadromous fishes 
in Canadian Arctic and marine waters. Given under scien- 
tific name, authority, and description date, are the Common 
Name, Provinces/Territories of occurrence, Ecozone, Distribu- 
tion (ocean and extralimital), Numbers, Size (maximum), and 
Biology (depth, food, commercial importance). An extralimitial 
list gives 83 species occurring in adjacent areas that might 
eventually be recorded in Canadian Arctic waters annotated 
as eastern (eastern Davis Strait, southwest Greenland) and 
western (western Beaufort Sea of Alaska from Point Barrow 


Marine Turtle Newsletter (103) 


January 2004. 32 pages: ARTICLES: Behavior of Green Sea 
Turtles in the Presence and Absence of Recreational Snork- 
ellers — Turtle Strandings in the Southern Eritrean Red Sea 
— First Confirmed Occurrence of Loggerhead Turtles in 
Peru — A Harness for Attachment of Satellite Transmitters 
on Flatback Turtles — Notes: Nesting of the Hawksbill Turtle 
in Shidvar Island, Hormozgan Provance, Iran — MEETING 
REPORTS — OBITUARIES (Henry H. Hildebrand 1922-2003; 
C. Robert Shoop 1935-2003) — ANNOUNCEMENTS — NEWS 
& LEGAL BRIEFS — RECENT PUBLICATIONS. 


eastwards). A brackish water list gives 36 species whose life 
is usually spent entirely in fresh waters but which may enter 
estuaries and seashore pools in the Canadian Arctic. There 
are 3 tables (species by province and territory, by ecozones, 
and by distribution (Alaska, Atlantic, Eurasia)); 2 maps (Can- 
adian Arctic waters and ecozones with this area); and two 
appendicies: one giving an alphabetical listing of scientific 
names of families with common names in English, French, 
Inuktitut and Inuvialuktun and one giving an alphabetical 
listing of species by scientific name with notation of new to 
Arctic, new to Canada, and page number in text. 

Copies of this report are available from Fisheries and 
Oceans Canada Central and Arctic Region, Winnipeg, Mani- 
toba R3T 2N6 Canada: e-mail reistj@dfo-mpo.ge.ca [Cata- 
logue number Fs 97-4/26/2674E, ISSN 0706-6473]. 


The Marine Turtle Newsletter is edited by Brendan J. 
Godley and Annette C. Broderick, Marine Turtle Research 
Group, School of Biological Sciences, University of Exeter, 
Exeter EX4 4PS United Kingdom; e-mail MTN @seaturtle. 
org; Fax +44 1392 263700. Subscriptions and donations 
towards the production of both the MTN and its Spanish 
edition NTM [Noticiero de Tortugas Marinas] should be 
made online at <http://www.seaturtle.org/mnt/> or c/o 
SEATURTLE.ORG, 11400 Classical Lane, Silver_Spring, 
Maryland 20901 USA. 


Ontario Natural Heritage Information Centre Science and Information Newsletter 9(1) 


Winter 2004 


Contents of this 20 page issue: FEATURE ARTICLES: Decade 
of Achievement (NHIC 10" anniversary celebrated 9 Decem- 
ber 2003) — A Decade of Ontario Botanical Dsicoveries; 
2003 PROGRAM HiIGLicuts: NHIC Staff Participate in James 
Bay Coastal Vegetation Study — Notes on the Vegetation 
and Flora of Flooding River Shores — A Profile of Natural 
Areas Recently Added to the Natural Areas Database; 2003 
ProJEcT HIGHLIGHTS: NHIC Conducts Turtle Surveys — 
NHIC Coordinates Odonata and Reptile Surveys — Update 
on the Great Lakes Conservation Blueprint Projects — 
Identifying Key Natural Areas and Linkages in Southern 
Ontario — NHIC’s Southern Region Element Occurrence 
Prioritization Project; NEws AND NorTEs: Species at Risk: A 


Diversity of Designations — What’s in a Name? Changes to 
the Provincial Amphibian and Reptile Lists — NHIC Attends 
Pelee Island Events — NatureServe Canada and Parks Agency 
sign MOU — NatureServe Participates with CWS on Spe- 
cies at Risk Information Needs Analysis — Additions to the 
COSEWIC list — Minister visits NHIC; Book REvVIEws: 
NHIC STAFF INFORMATION. 

Mailing address for Natural Heritage Information Centre, 
Ontario Ministry of Natural Resources, 300 Water Street, 
2nd Floor, North Tower, P.O. Box 7000, Peterborough, On- 
tario K9J 8M5, Canada; www.mnr.gov.on.ca/MNR/nhic.cfm. 
Web page: http://www.mnr.gov.on.ca/MNR/nhic.html 


2004 NEWS AND COMMENT 157 


Erratum Canadian Field-Naturalist 117(1) 


Ballard, Warren B., Matthew A. Cronin, Martin D The abbreviation for concentrations in micrograms per 
5 9 . 5 a . x z ; 
Robards, and William A. Stubblefield. 2003. Heavy gram of dry weight should be corrected in two places. 
, ‘ ; . x 0 as J. c rd ae 5 te pS) =f; k Pa ; Ore 
metal concentrations in Arctic Foxes, Alopex lagopus, in On page 120 “mg/g” left column line 18, and right 


the Prudhoe Oil Field, Alaska. Canadian Field-Naturalist column line 21, should be “g/g”. 
117(2): 119-121. 


Erratum Canadian Field-Naturalist 117(2) 


Lindquist, E. S., C. F. Aquadro, D. McClearn, and K J. 117(2): 184-189. 
McGowan. 2003. Field identification of the mice On page 4, Figure 1, 1A was repeated for 1B. The correct 
Peromyscus leucopus noveboracensis and P. maniculatus 1B is shown below with IA. 
gracilis in central New York. Canadian Field-Naturalist 


Distance from P. leucopus 
Distance from P. leucopus 


O 4 2 38 4 5 6 
Distance from P. maniculatus 


Distance from P. maniculatus 


FiGurRE 1. Classification of P. |. noveboracensis and P. m. gracilis using the discriminant-function coefficients given in Table 
2. Figure la shows the classification of adults only. Figure 1b shows the classification of adults and juveniles. Open 
and closed symbols denote P. m. gracilis and P. |. noveboracensis, respectively. In Figure 1b, P. m. gracilis is indicated 
by open circles and squares (adult and juvenile, respectively), and P. |. noveboracensis by closed triangles and stars 
(adult and juvenile, respectively). 


Advice for Contributors to The Canadian Field-Naturalist 


Content 

The Canadian Field-Naturalist is a medium for the publi- 
cation of scientific papers by amateur and professional natu- 
ralists or field-biologists reporting observations and results 
of investigations in any field of natural history provided that 
they are original, significant, and relevant to Canada. All read- 
ers and other potential contributors are invited to submit for 
consideration their manuscripts meeting these criteria. The 
journal also publishes natural history news and comment items 
if judged by the Editor to be of interest to readers and sub- 
scribers, and book reviews. Please correspond with the Book 
Review Editor concerning suitability of manuscripts for this 
section. For further information consult: A Publication Policy 
for the Ottawa Field-Naturalists’ Club, 1983. The Canadian 
Field-Naturalist 97(2): 231-234. Potential contributors who 
are neither members of The Ottawa Field-Naturalists’ Club 
nor subscribers to The Canadian Field-Naturalist are encour- 
aged to support the journal by becoming either members or 
subscribers. 


Manuscripts 

Please submit, to the Editor, in either English or French, 
three complete manuscripts written in the journal style. 
The research reported should be original. It is recommended 
that authors ask qualified persons to appraise the paper before 
itis submitted. All authors should have read and approved it. 
Institutional or contract approval for the publication of the data 
must have been obtained by the authors. Also authors are ex- 
pected to have complied with all pertinent legislation regard- 
ing the study, disturbance, or collection of animals, plants or 
minerals. The place where voucher specimens have been de- 
posited, and their catalogue numbers, should be given. Lati- 
tude and longitude should be included for all individual local- 
ities where collections or observations have been made. 

Print the manuscript on standard-size paper, doublespace 
throughout, leave generous margins to allow for copy mark- 
ing, and number each page. For Articles and Notes provide 
a bibliographic strip, an abstract and a list of key words. 
Generally, words should not be abbreviated but use SI symbols 
for units of measure. The names of authors of scientific names 
should be omitted except in taxonomic manuscripts or other 
papers involving nomenclatural problems. “Standard” com- 
mon names (with initial letters capitalized) should be used at 
least once for all species of higher animals and plants; all 
should also be identified by scientific name. 

The names of journals in the Literature Cited should be 
written out in full. Unpublished reports and web documents 
should not be cited here but placed in the text or in a sepa- 
rate Documents Cited section. List the captions for figures 
numbered in arabic numerals and typed together on a separate 
page. Present the tables each titled, numbered consecutively 
in arabic numerals, and placed on a separate page. Mark in 
the margin of the text the places for the figures and tables. 


Check recent issues (particularly Literature Cited) for 
journal format. Either “British” or “American” spellings are 
acceptable in English but should be consistent within one 
manuscript. The Oxford English Dictionary, Webster’s 
New International Dictionary and le Grand Larousse 
Encyclopédique are the authorities for spelling. 


Illustrations 

Photographs should have a glossy finish and show sharp 
contrasts. Electronic versions should be high resolution. Photo- 
graphic reproduction of line drawings, no larger than a 
standard page, are preferable to large originals. Prepare line 
drawings with India ink on good quality paper and letter (don’t 
type) descriptive matter. Write author’s name, title of paper, and 
figure number on the lower left corner or on the back of each 
illustration. 


Reviewing Policy 

Manuscripts submitted to The Canadian Field-Naturalist 
are normally sent for evaluation to an Associate Editor (who 
reviews it or asks another qualified person to do so), and at 
least one other reviewer, who is a specialist in the field, cho- 
sen by the Editor. Authors are encouraged to suggest names 
of suitable referees. Reviewers are asked to give a general 
appraisal of the manuscript followed by specific comments 
and constructive recommendations. Almost all manuscripts 
accepted for publication have undergone revision—sometimes 
extensive revision and reappraisal. The Editor makes the 
final decision on whether a manuscript is acceptable for pub- 
lication, and in so doing aims to maintain the scientific quality, 
content, overall high standards and consistency of style, of 
the joumal. 


Special Charges — Please take note 

Authors must share in the cost of publication by paying 
$80 for each page, plus $15 for each illustration (any size up 
to a full page), and up to $80 per page for tables (depending 
on size). Authors may also be charged for their changes in 
proofs. Reproduction of color photos is extremely expensive; 
price quotations may be obtained from the Business Manager. 
Reprint order forms are included when galley proofs are sent 
to authors. If grant or institutional funds are not available, club 
members and subscribers may apply for a waiver of charges 
for the first five pages. 

Limited joumal funds are available to help offset publi- 
cation charges to authors with minimal financial resources. 
Requests for financial assistance should be made to the Busi- 
ness Manager when the manuscript is accepted. 


Reprints 
An order form for the purchase of repents will accompany 
the galley proofs sent to the authors. 


FRANCIS R. Cook, Editor 
13078 Land O’Nod Road 
RR 3 North Augusta, Ontario KOG IRO Canada 


158 


TABLE OF CONTENTS (concluded) Volume 118 Number 1 


Record high wolf, Canis lupus, pack density L. DAVID MECH AND SHAWN TRACY 


Extraordinary size and survival of American Black Duck, Anas rubripes, broods 
JERRY R. LONGCORE AND DANIEL G. MCAULEY 


Observations of interactions between Puma, Puma concolor, and introduced 
European Red Deer, Cervus elaphus, in Patagonia WERNER T. FLUECK 


Premiéres mentions de la Couleuvre mince, Thamnophis sauritus septentrionalis, au Québec 
JEAN-FRANCOIS DESROCHES ET RICHARD LAPARE 


Book Reviews 

ZooLoGy: Amphibian Decline: An Integrated Analysis of Multiple Stressor Effects — Birds of Africa: 
From of Seabirds to Seedeaters — The Firefly Encyclopedia of Birds — Birds of Belize — Herpetology: 
Third Edition — Lizards: Windows to the Evolution of Diversity — Raptors of Western North America 
— Raptors of Eastern North America — Turtles and Tortoises — What Good are Bugs? Insects in the 
Web of Life — Belugas in the North Atlantic and Russian Arctic — The Firefly Encyclopedia of Insects 
and Spiders 


Botany: Biotic Forest Communities of Ontario — Les champignons des arbres de |’est de l’ Amerique du 
Nord — Manual of Vascular Plants of Northeastern United States and Adjacent Canada: Second Edition 
— The Wild Orchids of North America, North of Mexico — Trees of the Carolinian Forest: A Guide to 
Species, Their Ecology and Uses 


ENVIRONMENT: Good News for a Change: How Everyday People are Helping the Planet — Whose Bird? 
— Forest Dynamics and Disturbance: Studies from Temperate Evergreen-Deciduous Forests — 
Snowshoes & Spotted Dick: Letters from a Wilderness Dweller— Natural Grace: The Charm, Wonder, 
& Lessons of Pacific Northwest Animals & Plants — Tales from the Underground: A Natural History 
of Subterranean Life 


NEw TITLES 


News and Comment 

Point Pelee Natural History News 3(4) — COSEWIC Assessment Results November 2003 — Froglog (61) — 
The Boreal Dip Net 8(1) — Annotated List of the Arctic Marine Fishes of Canada — Marine Turtle News- 
letter (103) — Ontario Natural Heritage Information Centre Science and Information Newsletter 9(1) 
Winter 2004 — Erratum: The Canadian Field-Naturalist 117(1) — Erratum:The Canadian Field- 
Naturalist 117(2) 


Advise to Contributors 


Mailing date of the previous issue 117(4): 16 September 2004 


2004 


PA 


129 


138 


146 


149 
153 


THE CANADIAN FIELD-NATURALIST Volume 118 Number 1 200 
——_—— Ore NET EE UUE 


125 Years’ Anniversary: 


The Ottawa Field-Naturalists’ Club: origins and history DANIEL F. BRUNTON l 
Articles 
Continuing environmental change — an example from Nova Scotia EDMUND S. TELFER 39) 


Habitat segregation among songbirds using the southern boreal 
mixedwood forest ENID E. CUMMING 45 


Effects of mid-winter snow depth on stand selection by Wolverines, Gulo gulo luscus, 
in the boreal forest JONATHAN D. WRIGHT AND JESSICA ERNST 56 


Wolverine, Gulo gulo luscus, resting sites and caching behavior in the boreal forest 
JONATHAN D. WRIGHT AND JESSICA ERNST 61 


Seasonal home ranges of Raccoons, Procyon lotor, using a common feeding site in rural eastern 
Ontario: rabies management implications SARAH C. TOTTON, RICHARD C. ROSATTE, 
ROWLAND R. TINLINE, AND LAuRA L. BIGLER 65 


Status of marine turtles in British Columbia waters: a reassessment 
DONALD F. MCALPINE, STAN A. ORCHARD, 


KELLY A. SENDALL, AND RON PALM 2 
Introduced marine species in the Haida Gwaii (Queen Charlotte Islands) region, 
British Columbia N. A. SLOAN AND P. M. BARTIER Vil 
Ruby-throated Hummingbird, Archilochus colubris, entanglements in burdock 
(Arctium spp.) at Delta Marsh, Manitoba HEATHER L. HINAM, SPENCER G. SEALY, 
AND Topp J. UNDERWOOD 85 
Multiple mating results in multiple paternity in Richardson’s Ground Squirrels, 
Spermophilus richardsonii JAMES F. HARE, GLENDA Topp, AND WENDY A. UNTEREINER 90 
Demographic patterns and limitations of wolves, Canis lupus, in and near Pukaskwa 
National Park, Ontario S. ANNE FORSHNER, PAUL C. PAQUET, FRANK G. M. BURROWS, 
GRAHAM K. NEALE, AND KEITH D. WADE 95 


Estimation of seed bank and seed viability of the Gulf of Saint Lawrence Aster, 
Symphyotrichum laurentianum (Fernald) Nesom — Joni FE. KEMP AND CHRISTIAN R. LACROIX 105 


Consumption of shrews, Sorex spp., by Arctic Grayling, 
Thymallus arcticus JONATHAN W. Moore AND G. J. KENAGY Hi 


Unusual behavior by Bison, Bison bison, toward Elk, Cervus elaphus, and Wolves, Canis lupus 
L. Davip MEcH, RICK T. MCINTYRE, AND DOUGLAS W. SMITH 115 


r 


Notes 


Morphology of female Woodland Caribou, Rangifer tarandus caribou, 
in Saskatchewan W. JAMES RETTIE 119 


Occurrence of parasitoid wasps in the egg sacs of Pardosa moesta and Pardosa sternalis 
(Araneae, Lycosidae) in southern Idaho LisA M. COBB AND VINCENT A. CoBB 12D 


The Northern True Katydid, Pterophylla camellifolia (Insecta: Orthroptera: Pseudophyllidae), 
at Ottawa, Ontario STEPHEN J. DARBYSHIRE 124 


(continued on inside back cover) 


ISSN 0008-3550 


The CANADIAN 
— FIELD-NATURALIST 


Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada 


Volume 118, Number 2 April-June 2004 


The Ottawa Field-Naturalists’ Club 


FOUNDED IN 1879 


Patrons 
Her Excellency The Right Honourable Adrienne Clarkson, C.C., C.M.M., C.D. 
Governor General of Canada 
His Excellency John Ralston Saul, C.C. 


The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; te 
encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields 
as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environments) 
of high quality for living things. 


Honorary Members 


Edward L. Bousfield Bruce Di Labio John A. Livingston E. Franklin Pope 
Donald M. Britton R. Yorke Edwards Stewart D. MacDonald William O. Pruitt,, Jr. 1 
Irwin M. Brodo Anthony J. Erskine Hue N. MacKenzie Joyce and Allan Reddoch | 
William J. Cody John M. Gillett Theodore Mosquin Mary E. Stuart 
Francis R. Cook C. Stuart Houston Eugene G. Munroe John B. Theberge 
Ellaine Dickson George F. Ledingham Robert W. Nero Sheila Thomson 
2004 Council 

President: Mike Murphy Ronald E. Bedford Barbara Gaertner Stanley Rosenbaum 
Vice-President: Gillian Marston Fenja Brodo Diane Lepage Louise Schwartz 
Recording Secretary: Susan Laurie-Bourque John Cameron Christina Lewis David Smythe 
Treasurer: Frank Pope William J. Cody Karen McLachalan Hamilton Henry Steger 
Past President: Gary McNulty Kathy Conlan David Hobden Chris Traynor 

Francis R. Cook Cendrine Huemer 


To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate 
P.O. Ottawa, Canada K1Z 1A2, or e-mail: ofnc @achilles.net. 
For information on Club activities telephone (613) 722-3050 or check http//www.achilles.net/ofnc/index.htm 


The Canadian Field-Naturalist 


The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this 
journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. 


We acknowledge the financial support of the Government of Canada toward our mailing cost through the Publication Assistance 
Program (PAP), Heritage number 09477. 


Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KOG IRO; (613) 269-3211; e-mail: feook @achilles.net 
Copy Editor: Elizabeth Morton 

Business Manager: William J. Cody, P.O. Box 35069, Westgate P.O. Ottawa, Canada KIZ 1A2; (613) 759-1374 
Book Review Editor: Roy John, 2193 Emard Crescent, Ottawa, Ontario K1J 6K5, e-mail: roy.john@pwegsc.ge.ca 


Associate Editors: Robert R. Anderson Paul M. Catling David Nagorsen 
Charles D. Bird Brian W. Coad Donald F. McAlpine 
Robert R. Campbell Anthony J. Erskine William O. Pruitt, Jr. 


Chairman, Publications Committee: Ronald E. Bedford 


All manuscripts intended for publication except Book Reviews should be addressed to the Editor and sent by postal 
mail. Book-review correspondence should be sent by e-mail or postal mail to Roy John, Book-review Editor. 


Subscriptions and Membership 

Subscription rates for individuals are $28 per calendar year. Libraries and other institutions may subscribe at the rate of $45 per 
year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $28 (individual) $30 (family) $50 (sustaining) and 
$500 (life) includes a subscription to The Canadian Field-Naturalist. All foreign subscribers and members (including USA) must 
add an additional $5.00 to cover postage. The club regional journal, Trail & Landscape, covers the Ottawa District and Local Club 
events. It is mailed to Ottawa area members, and available to those outside Ottawa on request. It is available to Libraries at $28 per — 
year. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be mailed to: © 
The Ottawa Field-Naturalists Club, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. Canada Post Publications Mail 
Agreement number 40012317. Return Postage Guaranteed. Date of this issue: April—June 2004 (April 2005). 


Cover: Brown Bear, Ursus arcts, watches Gray Wolf, Canis lupus, in the Katmai National Park and Preserve in southwest Alaska. 
See paper by Smith, Partridge, and Shoen pages 247-250 (Kent Fredrikson photograph). 


The Canadian Field-Naturalist  Uxiijeees2y 


Volume 118, Number 2 April-June 2004 


Morphology and Population Characteristics of Vancouver Island Cougars, 
Puma concolor vancouverensis 


STEVEN F. WILSON!, APRYL HAHN, AARON GLADDERS, KAREN M. L. Gou, and DAvip M. SHACKLETON 


Wildlife Research Group, Agroecology, Faculty of Agricultural Sciences, University of British Columbia, 270-2357 Main 
Mall, Vancouver, British Columbia V6T 1Z4 Canada 
'Present address: EcoLogic Research, 406 Hemlock Avenue, Gabriola Island, British Columbia VOR 1X1 Canada 


Wilson, Steven F., Apryl Hahn, Aaron Gladders, Karen M. L. Goh, and David M. Shackleton. 2004. Morphology and population 
characteristics of Vancouver Island Cougars, Puma concolor vancouverensis. Canadian Field-Naturalist 118(2): 159-163. 


Cougars are a management concern on Vancouver Island because they are a top predator and because there have been frequent 
attacks on humans on the island. However, little is known about Cougar ecology in the Pacific Northwest of North America. 
We studied Cougar morphology and population characteristics as part of a larger study in two areas on Vancouver Island. We 
derived a multivariate measure of body size to describe changes with age and sex. Body size was similar in the two study 
areas. Survival rates for adult females were higher than those reported elsewhere; however, hunters avoided shooting females 
in general, and radio-collared Cougars in particular. Litter size at first detection was lower than reported in many other 
studies and may be related to food availability. 


Key Words: Cougars, Puma concolor vancouverensis, morphology, survival, mortality, natality, Vancouver Island, British 


Columbia. 


The Mountain Lion, or Cougar (Puma concolor van- 
couverensis), 1s one of the two main large predatory 
carnivores in the forests of Vancouver Island, British 
Columbia, Canada. It also has a long history of inter- 
actions with humans as a nuisance (livestock predation) 
and as a game species (Beier 1991). The incidence of 
Cougar attacks on humans in North America is also 
highest on the island. However, little is known about 
the ecology of Cougar in this part of its range, mainly 
because this subspecies inhabits dense forests typical 
of coastal British Columbia. We examined Cougar mor- 
phology and population characteristics in two study 
areas on Vancouver Island to investigate the effects of 
age, sex, and location on body size, and to determine 
survival rates and reproductive characteristics. 


Methods 

Research was conducted on a 700-km? area near 
Northwest Bay (NWB), British Columbia, and on a 
1000-km? area centred on the Adam and Eve River 
valleys (AE) approximately 300 km northwest of NWB 
(Figure 1). Principal habitats at NWB consisted of 
different seral stages of Douglas Fir (Pseudotsuga men- 
ziesii), Western Hemlock (Tsuga heterophylla), and 
Western Redcedar (Thuja plicata) forests. The AE 
study area was more rugged and dominated by exten- 
sive stands of unlogged forest, particularly at higher 


elevations. Habitats at lower elevations were dominated 
by Western Hemlock and Western Redcedar while spe- 
cies at higher elevations included Mountain Hemlock 
(Tsuga mertensiana) and Yellow Cedar (Chamaecy- 
paris nootkatensis). Understory vegetation on both 
study areas was typically dense and consisted of Salal 
(Gaultheria shallon), ferns (several genera) and Vac- 
cinium species communities. Both study areas had 
cool summers and mild winters, with temperatures of 
0-10°C for 4-6 months/year and annual precipitation 
of 1700-5000 mm/year (Meidinger and Pojar 1991). 
AE was cooler and wetter, with a greater proportion 
of precipitation falling as snow. 

Cougars were captured between 4 March 1991 and 
1 September 1996 at NWB and from 18 April 1997 to 
12 September 1998 at AE. Local houndsmen were 
contracted to track and tree Cougars. Treed Cougars 
that could safely be immobilized were darted with a 
mixture of ketamine hydrochloride (3 mg/kg estimat- 
ed total weight) and medetomidine hydrochloride (0.1 
mg/kg). Cougars were sexed, measured (see below), 
and aged according to tooth replacement and wear, and 
by gum recession (Ashman et al. 1983; Laundré et al. 
2000). Adult Cougars were fitted with VHF radio col- 
lars (Telonics, Inc., Mesa, AZ). Immobilizations were 
reversed with atipamezole (0.15 mg/kg). Morphologi- 
cal data from hunter-killed Cougars in and near AE 
were also included in our analyses. 


159 


160 


126° W 124° W 


50° N 


Bay (NWB) 


FiGureE |. Location of cougar study areas on Vancouver 
Island, British Columbia, Canada. 


Morphological measurements taken included: total 
weight (using a 100 kg spring balance to the nearest 
1 kg), neck circumference at the base of the skull, chest 
circumference immediately behind the front legs, 
body length from nose to base of tail, tail length from 
base to tip of the last vertebrae (all to the nearest cm 
with a cloth tape measure), canine length from 
gumline to tip, and front and rear pad widths (to the 
nearest mm with a cloth tape measure; AE only). 

We used principal component analysis (PCA) to 
derive a single variable to represent body size, based 
on the correlation matrix of morphological variables 
(Statistica 1995). We used this body size measure in- 
stead of total weight to describe the size of Cougars 
because it was less condition-dependent that using 
total weight alone. Where Cougars were captured >1 
time, we included data from only the most recent 
capture. We used an analysis of covariance to test for 
differences in body size (log-transformed) between 
sex and study area, using age as a covariate. We also 
calculated means (+2 SE) for each morphological 
variable (by sex), but excluded Cougars <2 years old 
to minimize skews in distributions caused by the 
smaller body sizes of juvenile Cougars. 

We calculated survival estimates with the staggered 
entry design of Pollock et al. (1989), which is based 
on the Kaplan-Meier product limit estimator (Kaplan 
and Meier 1958). This method estimates annual sur- 
vival rate as the product of weekly survival rates (1 — 
d/r), where d is the number of animals that die and r 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


is the number of animals “at risk” in a weekly period. 
Cougars carrying functioning radio collars comprised 
the “‘at risk” sample for each week. The design was 
“staggered entry” because not all Cougars carried col- 
lars at the same time. Cougars were added to the week- 
ly sample as they were collared, and were removed as 
they died, had their collars removed, or when their col- 
lars stopped functioning. Analysis started with the 
collaring of the first Cougars in the two study areas. 
Annual survival estimates were based on 52 consecu- 
tive, seven-day periods. 

There were sufficient data to analyse adult female 
Cougar survival; however, sample sizes were small, 
and the length of the study was relatively short in 
relation to the life expectancy of Cougars. Therefore, 
our estimates should be interpreted with caution. 

We report the reproductive characteristics of col- 
lared females anecdotally because data were insuffi- 
cient to calculate population growth rates; specifical- 
ly, we had few data on birth intervals and survival to 
maturity. We report litter sizes (42 SE) when they were 
first seen, so litter sizes at birth may have been larger, 
although in at least three litters, kittens were seen when 
<10 days old (eyes not open). 


Results 

Thirty Cougars were measured at NWB (17 females 
and 13 males) and 26 at AE (15 females and 11 males). 

We used total weight, neck and chest circumfer- 
ence, and total length (body length + tail length) in 
the final PCA to derive a body size variable. All four 
variables loaded strongly and positively on the first 
axis, which explained 83% of the variation in the 
dataset. As a result, we used the first PCA axis as the 
index of body size (Table 1). We did not include canine 
length and body and tail lengths as separate variables 
in the analysis because doing so did not significantly 
increase the variation described in the first axis. 

Body size differed between males and females 
(P < 0.000), but not between study areas (P = 0.736; 
Figure 2). Differences between males and females were 
also evident in univariate means of the morphologi- 
cal measurements (Table 2). 

Mean survival rates were similar for collared Cou- 
gars on both study areas (Table 3). Known causes of 
female mortality at NWB included intraspecific kill- 
ings (2) and animal control (3). The cause of one fe- 
male mortality at NWB and both female mortalities at 
AE were unknown. Two mortalities of collared male 


TABLE |. Correlation between morphological variables and the first principal component axis (also known as “factor load- 
ings”). The axis (based on the correlation matrix) was used as an index to describe body size of Vancouver Island Cougars 


in subsequent analyses. 


Factor loadings of morphological variables 
Neck Chest 
0.889 0.927 


Total Weight 
0.994 


Body length 


Eigenvalue 
3.314 


% variation explained 


0.878 82.8 


2004 


WILSON, HAHN, GLADDERS, GOH, and SHACKLETON: VANCOUVER ISLAND COUGARS 16] 


Males: 
y = 1.45Ln(x) - 0.54 


R? = 0.70 


Females: e 
6 y = 0.52Ln(x) - 1.09 
R? = 0.34 


UF T 


0 2 4 


T li 


6 8 10 


Age (years) 


FiGure 2. Relationship between body size (derived from the principal component analysis on morphological measurements, 
see text) and age for male and female Cougars in the two Vancouver Island study areas. Logarithmic trend lines are 
presented for males (squares @) and females (diamonds @) , pooled by study area. The body size measurement was 
used because it was less condition-dependent than using total weight alone. 


Cougars were investigated at AE; one was shot by hunt- 
ers and the other was suspected to have been shot 
illegally. 

At NWB, mean litter size at first detection was 1.9 
+ 0.1 (n= 16). One female was known to have a litter 
of three kittens, and there were 12 other litters of two 
kittens and 3 of one kitten when first observed. Fe- 
males were known or suspected to give birth during all 
months from March-October. Consecutive litters were 
recorded for two females; each had litters 21 and 23 
months apart. 


Litter size at AE was 1.8 + 0.7 (n = 5). There was 
one litter of three, and two litters each of one and two 
kittens. Females gave birth in January, August, and 
September. No collared females at AE had more than 
one litter during the study period. 


Discussion 

Results presented in this paper are based on small 
absolute sample sizes; however, Cougars are rare and 
secretive, and inferences about their morphology and 
behaviour must often be based on few animals. The 


TABLE 2. Means and standard errors of morphological measurements taken on captured Vancouver Island Cougars. 


Males 
Measurement n Mean 
Total Weight (kg) 13 Ss 
Neck (cm) 12 39.5 
Chest (cm) 12 VS) 
Body length (cm) 13 128.2 
Tail Length (cm) 13 73.0 
Front pad Width (mm) 9 57.9 
Hind pad Width (mm) 9 50.4 
Canine Length (mm) 10 28.6 


Females 
2 SE n Mean 2 SE 
Si 22 39.1 Pel 
3.0 23 394 13} 
3.5 23 65.4 1.6 
6.6 23 120.9 Bill 
4.5 23 ORI 3} 
1.9 12 Syl I) 
33 12 37.9 9.1 
es) 23 26.8 2.0 


162 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE 3. Annual and mean survival rates of female Vancouver Island Cougars collared in two study areas: Northwest Bay 
(NWB) and Adam and Eve rivers (AE). Annual estimates were based on a staggered-entry design, starting with the first 
Cougars collared at NWB and AE in March 1991 and April 1997, respectively. 


Study Area Year n 


NWB 1991-1992 
1992-1993 
1993-1994 
1994-1995 
1995-1996 
AE 1997-1998 

1998-1999 

1999-2000 


ISSAP Sy Sap ays 


densities of Cougars in our study areas also suggest 
that significant proportions of the populations were 
captured (2.6-7.3 Cougars/100 km? at NWB and 1.4- 
2.0 Cougars/100 km? at AE; S. Wilson, unpublished 
data). Our sample sizes were similar to, or larger than, 
those in many comparable studies (e.g., n = 22, 
Hemker et al. 1984; n = 68, Ross and Jalkotzy 1992; 
n= 76, Lindzey et al. 1994; n = 34, Spreadbury et al. 
19962 n=13;, Prankhin;et ‘al. 1999: n= 21, Pierce;ct 
al. 2000). 

Few researchers have published Cougar morpholog- 
ical characteristics (c.f Kohlmann and Green 1999; 
Grigione et al. 2002); however, mean measurements 
of Vancouver Island Cougars in this study were smaller 
than those reported by Cowan and Guiguet (1965) for 
14 adult male (mean total length 241 cm, tail length 
89 cm, mean total weight 73 kg) and 7 adult female 
(mean total length 206 cm, tail length 79 cm, mean 
total weight 46 kg). Vancouver Island Cougars are 
slightly smaller than those found elsewhere in British 
Columbia (Cowan and Guiguet 1965). 

Survival estimates are critical for population man- 
agement, but few studies report them. Using the same 
method and similar sample sizes, Lindzey et al. (1988) 
reported mean survival rates (S = 0.731) for female 
Cougars in a largely unhunted population in southern 
Utah that were lower than those we calculated for 
Vancouver Island Cougars (mean for both study areas 
S = 0.885). Hunting is an important component of 
mortality in Cougar populations where hunting is al- 
lowed (Hemker et al. 1984; Logan et al. 1986; Ross 
and Jalkotzy 1992). The Vancouver Island population 
is no exception; however, we were in frequent contacts 
with hunters in our study areas and know they often 
treed radio-collared cougars but chose not to shoot 
them, even though we did not request this. This might 
have inflated the survival rates calculated for radio- 
collared Cougars. Researchers and managers should 
be aware of this bias when calculating survival esti- 
mates from similar radio telemetry studies where 
hunters “lend a hand” by not shooting study animals. 


Survival -2 SE +2 SE Mean 
1 
1 
0.75 0.47 1 
0.82 0.56 1 
0.88 0.64 1 0.89 
0.80 0.53 1 
1 
0.83 0.50 ff 0.88 


Animal control by British Columbia Wildlife Branch 
Control Officers was a significant source of human- 
caused mortality at NWB, where the human popula- 
tion density was higher than at AE. Intraspecific killing 
has been reported as an important source of mortality 
elsewhere (Lindzey et al. 1988; Spreadbury et al. 1996), 
but in our study it was detected only at NWB where 
Cougar population density was significantly higher than 
at AE. 

Litters in both of our study areas were smaller than 
those reported in southeastern British Columbia 
(x = 3.1; Spreadbury et al. 1996), southwestern Al- 
berta (x = 2.2; Ross and Jalkotzy 1992), Wyoming 
(x = 2.7; Logan et al. 1986), and southern Utah 
(x = 2.4; Lindzey et al. 1994). Other studies have 
recorded births in most months of the year (Ross and 
Jalkotzy 1992; Lindzey et al. 1994), with a peak in 
late summer and early fall (Lindzey et al. 1994). Our 
observations at AE were similar. The pattern was dif- 
ferent at NWB, with no births recorded in winter and 
no obvious peaks in births during the spring-fall period. 

Columbia Black-tailed Deer (Odocoileus hemionus 
columbianus) are the Vancouver Island Cougars’ pri- 
mary prey. Indices of deer abundance declined 55% 
from 1991-1996 at NWB and 38% from 1995-1999 at 
AE (British Columbia Ministry of Environment, Lands 
and Parks, unpublished data); therefore, small litter 
sizes among Vancouver Island Cougars may be a result 
of low food availability. Also, Black-tailed Deer are 
smaller than mainland Mule Deer (Odocoileus hemi- 
onus; Shackleton 1999). This highlights the importance 
of maintaining prey populations in management of 
Cougars on Vancouver Island. 


Acknowledgments 

This research was supported by Forest Renewal 
British Columbia, British Columbia Ministry of Envi- 
ronment, Lands, and Parks (MELP), Habitat Conserva- 
tion Trust Fund, Science Council of British Columbia, 
MacMillan Bloedel Limited, TimberWest Limited, 
Western Forest Products Limited, BC Hydro, Mountain 


2004 


Equipment Co-op, and the University of British Col- 
umbia. We thank M. DeLaronde, S. Foster, D. Koshow- 
ski, J. Leo, R. Ramcharita, H. Robinson, C. Wainwright, 
K. Wickert, and local houndsmen for assistance with 
fieldwork. For logistic support we also thank G. Brun- 
ham, K. Brunt, T. Hamilton, D. Janz, and H. Schwantje 
of MELP, as well as many staff of MacMillan Bloedel 
Limited in Nanaimo and Campbell River, British 
Columbia, and of TimberWest Limited in Nanaimo, 
B.C. S. Kohlmann, F. Scott and D. Nagorsen made 
helpful comments on the manuscript. This paper is 
dedicated to the memory of the late K. Atkinson, who 
conceived and supported the project from the outset. 


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Received 7 May 2001 
Accepted 5 November 2004 


Conservation Evaluation of Lemmon’s Holly Fern, Polystichum 
lemmonii, a Threatened Fern in Canada” 


GEORGE W. DoucLas!: 2 


‘Conservation Data Centre, British Columbia Ministry of Sustainable Resource Management, Terrestrial Information 
Branch, P.O. Box 9993 Stn Prov Govt, Victoria, British Columbia V8W 9R7 Canada 
> Current address: Douglas Ecological Consultants Ltd., 6230 North Road, Duncan, British Columbia V9L 6K9 Canada 


Douglas, George W. 2004. Conservation evaluation of Lemmon’s Holly Fern, Polystichum lemmonii; a threatened fern in 


Canada. Canadian Field-Naturalist 118(2): 164-168. 


In Canada, Lemmon’s Holly Fern, Polystichum lemmonii, is restricted to the Baldy Mountain area on the eastern side of the 
Okanagan River valley in south-central British Columbia. This population represents the northern limits of the species 
which ranges south through northern Idaho, Washington and Oregon to northern California. In British Columbia, P. lemmonii 
is associated with ultramafic rocky ridges within a montane forest at an elevation of 1900 m. The population in the Baldy 
Mountain area is relatively small, unprotected and potentially imperilled by mining exploration, forest road construction or 


wildfires. 


Key Words: Lemmon’s Holly Fern, Polystichum lemmonii, threatened, distribution, population size, British Columbia. 


Lemmon’s Holly Fern, Polystichum lemmonii 
Underw. [taxonomy and nomenclature follow Douglas 
et al. (1998a, b; 2000)], is a member of a cosmopolitan 
genus of over 175 species (Smith and Lemieux 1993; 
Wagner 1993). It is one of eight Polystichum species 
occurring in British Columbia (Ceska 2000) and nine 
occurring in Canada (Cody and Britton 1989; Wagner 
1993). Generally, American authors treated P. lem- 
monii as a synonym of P. mohrioides (Bory) C. Presl. 
until Wagner (1979) demonstrated that the North Am- 
erican plant was different from the South American 
plant. Polystichum lemmonii was first recorded in 
Canada by Cody and Britton (1984). 

Polystichum lemmonii 1s an evergreen, perennial, 
tufted fern arising from a short, stout rhizome (Figure 
1; Ceska 2000). The decumbent to ascending fronds 
are 10-40 cm long, 3-7 cm wide and 2-pinnate. The 
20-35 pinnae on each side of the rachis are ovate with 
rounded pinnules. The ultimate segments are entire 
or weakly toothed. The round sori are attached near 
the midvein with entire or minutely toothed indusia. 

In British Columbia, P. lemmonii may be confused 
with either Kruckeberg’s Holly Fern (P. kruckebergii) 
or Mountain Holly Fern (P. scopulinum). It may be 


* The field work for the Polystichum lemmonii project was 
funded by the British Columbia Conservation Data Centre. 
The results appear in the British Columbia Conservation 
Data Centre database and a rare plant manual (Douglas 
et al. 2002). This information formed the basis for a 
Committee on the Status of Endangered Wildlife in Canada 
status report (Douglas 2003*) and the subsequent assess- 
ment of threatened (COSEWIC 2003*). The present paper 
also includes more recent information that will be used 
in a National Recovery Strategy for P. lemmonii (Doug- 
las 2005). 


FiGurE 1. Mlustration of Polystichum lemmonii (line 
drawing from Ceska 2000). 


164 


2004 


———~ 
KAMLOOPS 


SPENCES BRIDGE 
ARMSTRONG 


Sw 


PRINCETON PENTICTON 


KEREMEOS | 


) BRITISH 
‘& . COLUMBIA 


FIGURE 2. The small square indicates the location of 
Polystichum lemmonii in British Columbia. 


DOUGLAS: LEMMON’S HOLLY FERN IN CANADA 165 


distinguished from the latter two species by the lack 
of spines on the teeth of the ultimate segments of the 
pinnae (Hitchcock et al. 1969; Wagner 1993; Ceska 
2000). 


North American and Provincial Ranges 

Polystichum lemmonii ranges from south-central 
British Columbia, sporadically south through Wash- 
ington and Oregon to northern California (Smith and 
Lemieux 1993; Wagner 1993). In Canada, it is known 
only from the Baldy Mountain area in the Okanagan 
River valley in south-central British Columbia (Figure 
2; Ceska 2000; Douglas et al. 2002). 


Habitat 

In western North America, Polystichum lemmonii 
occurs on sites where ferromagnesian or ultramafic 
rocks outcrop (Kruckeberg 1969; Wagner 1993). There 
are a number of ultramafic rock outcrops in western 
British Columbia but only the two small, adjacent 
ridges, at an elevation of 1900 m, in the Baldy Moun- 
tain area support the latter species (Figure 3). These 
dunite rock outcrop habitats are also characterized by 
shallow soils thus creating xeric microclimates that 
exclude many nearby species adapted to more mesic 
microclimates or non-ultramafic soils (Figure 4). These 
ridges, therefore, have a typically depauperate ultra- 
mafic flora and lack a tree cover in contrast to the 


FicureE 3. Aerial view of the ultramafic east ridge. Most Polystichum lemmonii plants occur on the north side 
of these east-west oriented ridges. The west ridge is slightly longer. 


166 


surrounding montane forests. The most prominent 
species on the ridge include Common Juniper (Juni- 
perus communis), Indian’s Dream (Aspidotis densa), 
Alpine Sandwort (Minuartia obtusiloba) and Yarrow 
(Achillea millefolium). 


Biology 

There is a limited amount of information on the 
biology and ecology of Polystichum lemmonii. The 
ultramafic habitat, however, has been well-studied by 
Kruckeberg (1969). Plants of ultramafic substrates 
are adapted to tolerate low levels of calcium, nitro- 
gen, phosphorus and molybdenum and high levels of 
magnesium, chromium and nickel (Kruckeberg 1969). 

Wagner (1979) has provided some genetic informa- 
tion. P. lemmonii is a tetraploid (2n = 82) and thought 
to be one of the parents of both P. kruckebergii and P. 
scopulinum. 

Sporophytes of P. lemmonii, as with most evergreen 
fern species, often retain significant numbers of mature 
spores over the winter that are then released the fol- 
lowing spring (Farrar 1976). PR. lemmonii also grows 
vegetatively by subterranean rhizome elongation 
often resulting in large clumps of clones. Because of 
the dry site conditions, which are not ideal for spore 
germination or gamete fertilization, most reproduction 
is probably by rhizome elongation (Walker 1979). 
Prothalli were not observed at the site. 

Long-distance dispersal of spores of P. lemmonii is 
evident by the distance to the nearest locations of the 
species in the adjacent state of Washington in the 
United States. P. lemmonii occurs in the Twin Sisters 
Range of Washington and in the Wenatchee Mountains 
(Kruckeberg 1969), a distance of about 205 km to the 
southwest and 225 km to the south of Baldy Moun- 
tain, respectively. The Tulameen River ultramafic site, 
where both P. kruckebergii and P. scopulinum occur 
(Kruckeberg 1969; Douglas and Labrecque 2003%*), 
does not contain P. lemmonii even though the Tula- 
meen River site is halfway between Baldy Mountain 
and the Twin Sisters Range. Reproduction of P. lem- 
moni at the Baldy Mountain area is evident since about 
30 percent of the 853 plants counted were relatively 
young (plant tufts less than four cm wide). 


Population Attributes 

The population of P. lemmonii occurs on two adja- 
cent, rocky ridges. The ridges, which are about 280 and 
200 m long by 50 m wide, are separated by a distance 
of 160 m. A 2001 count of all plants in the popula- 
tion by the author revealed a total of 853 plants over 
0.72 ha. A collection by D. M. Britton in 1987 (at 
Department of Agriculture, Ottawa) mentions a popu- 
lation size of “perhaps a thousand plants”. This would 
indicate that the population has remained relatively 
stable for at least 15 years. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Provincial, National and Global Ranks 

The British Columbia Conservation Data Centre 
has ranked this species as S1 and placed it on the 
British Columbia Ministry of Sustainable Resource 
Management Red-list (Douglas et al. 2002). This is 
the most critical category for imperilled rare native 
vascular plants in British Columbia. A rank of S1 is 
considered “critically imperilled because of extreme 
rarity (5 or fewer occurrences or very few remaining 
individuals) or because of some factors making it 
especially vulnerable to extirpation or extinction” 
(Douglas et al. 2002). Since the species is restricted 
to British Columbia, the National rank is N1. Globally, 
Polystichum lemmonii is ranked G4 and is frequent to 
common in its range and apparently secure. 


Threats and Protection 

The most direct threat to Polystichum lemmonii is 
mining exploration. At the present time the entire 
area is occupied by active mining claims and explora- 
tion could occur with short notice. Additional threats 
include the potential use of the rock outcrop for forest 
road construction and the possibility of intense wild- 
fires. Extremely high forest fuel loads in the the area 
may lead to wildfires similar to that experienced in 
the region in 2003. Introduced species are of no con- 
cern at this site due to the ultramafic properties of the 
soils. 

The population in the Baldy Mountain area is on 
public land but is not part of a protected area. It is 
conceivable that this area could qualify as a Wildlife 
Habitat Area but this status has yet to be proposed. 
Polystichum lemmonii could be a candidate species 
for protection under the provincial Wildlife Amend- 
ment Act as it is currently Red-listed by the British 
Columbia Conservation Data Centre. 


Evaluation 

The British Columbia Conservation Data Centre 
considers Polystichum lemmonii to be threatened/en- 
dangered in British Columbia (Douglas et al. 2002) 
and the Committee on the Status of Endangered Wild- 
life in Canada has assessed the species as threatened 
(COSEWIC 2003). Just over 850 plants are known 
from a single site in Canada at the Baldy Mountain 
area of south-central British Columbia. The prognosis 
for this species is not good since ultramafic rock out- 
crops often attract mineral exploration and the entire 
area is covered by active mineral claims. These rock 
outcrops could also be of potential use as a quarry for 
road building materials. The extremely high forest fuel 
loads in the adjacent area could also support wildfires. 
Establishment of a Wildlife Habitat Area and removal 
of the active mineral claims would remove the major 
threats at the site. 


2004 


DOUGLAS: LEMMON’S HOLLY FERN IN CANADA 


167 


FicureE 4. Polystichum lemmonii plants are conspicuous among the low vegetation on the ultramafic soils. 


Acknowledgments 

I would like to thank Kathy Paige for her partici- 
pation in the fieldwork and Louise Bacon and Jenifer 
Penny for office assistance. Graham Nixon, British 
Columbia Geological Survey, kindly provided infor- 
mation on ultramafic rock types. 


Documents Cited (marked * in text) 

COSEWIC. 2003. COSEWIC assessment and status report 
on Lemmon’s holly fern Polystichum lemmonii in Canada. 
Committee on the Status of Endangered Wildlife in Can- 
ada. www.cosewic.gc.ca, Ottawa, Ontario. 13 pages 

Douglas, G. W. 2003. COSEWIC Status report on Lemmon’s 
holly fern Polystichum lemmonii in Canada, in COSEWIC 
assessment and status report on Lemmon’s holly fern 
Polystichum lemmonii in Canada. Committee on the Status 
of Endangered Wildlife in Canada. www.cosewic.gc.ca, 
Ottawa, Ontario. 13 pages. 

Douglas, G. W., and J. Labrecque. 2003. COSEWIC Status 
report on Mountain Holly Fern or Polystic des Rochers, 
Polystichum scopulinum in Canada. Unpublished report. 
Ottawa, Ontario. 20 pages. 

Douglas, G. W. 2005. National Recovery Strategy for Lem- 
mon’s Holly Fern (Polystichum lemmonii). In preparation. 
Ottawa, Ontario. 


Literature Cited 
Ceska, A. 2000. Pteridophytes. Pages 260-343 in Illustrated 
flora of British Columbia. Volume 5. Dicotyledons (Salica- 


ceae through Zygophyllaceae) and Pteridophytes. Edited 
by G. W. Douglas, D. Meidinger and J. Pojar. British 
Columbia Ministry of Environment, Lands and Parks and 
British Columbia Ministry of Forests. Victoria, British 
Columbia. 427 pages. 

Cody, W. J., and D. M. Britton. 1984. Polystichum lemmonii, 
a rock shield-fern new to British Columbia and Canada. 
Canadian Field-Naturalist 98: 375. 

Cody, W. J., and D. M. Britton. 1989. Ferns and fern allies 
of Canada. Agriculture Canada Research Branch, Publica- 
tion 1829/E. Ottawa, Ontario. 430 pages. 

Douglas, G. W., D. Meidinger, and J. L. Penny. 2002. Rare 
native vascular plants of British Columbia. Second edition. 
Province of British Columbia. Victoria, British Columbia. 
359 pages. 

Douglas, G. W., D. Meidinger, and J. Pojar. 2000. Illus- 
trated flora of British Columbia. Volume 5. Dicotyledons 
(Salicaceae through Zygophyllaceae) and Pteridophytes. 
British Columbia Ministry of Environment, Lands and 
Parks and British Columbia Ministry of Forests. Victoria, 
British Columbia. 389 pages. 

Douglas, G. W., G. B. Straley, and D. Meidinger. 1998a. 
Illustrated flora of British Columbia. Volume 1. Gymno- 
sperms and Dicotyledons (Aceraceae through Asteraceae). 
British Columbia Ministry of Environment, Lands and 
Parks and British Columbia Ministry of Forests. Victoria, 
British Columbia. 436 pages. 

Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar. 
1998b. Illustrated flora of British Columbia. Volume 2. 
Dicotyledons (Balsaminaceae through Cuscutaceae). Bri- 


168 


tish Columbia Ministry of Environment, Lands and Parks 
and Ministry of Forests. Victoria, British Columbia. 401 
pages. 

Farrar, D. L. 1976. Spore retention and release from over- 
wintering fronds. American Fern Journal 66: 49-52. 

Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W. 
Thompson. 1969. Vascular plants of the Pacific Northwest. 
Part 1. Vascular Cryptogams, Gymnosperms and Mono- 
cotyledons. University Washington Press, Seattle, Wash- 
ington. 914 pages. 

Kruckeberg, A. R. 1969. Plant life on serpentine and other 
ferromagnesian rocks in northwestern North America. 
Syesis 2: 14-114. 

Smith, A. R., and T. Lemieux. 1993. Dryopteridaceae. Pages 
91-94 in The Jepson manual: Higher plants of California. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Edited by J. C. Hickman. University of California Press. 
Berkeley, California. 1400 pages. 
Wagner, D. H. 1979. Systematics of Polystichum in western 
North America north of Mexico. Pteridologia 1: 1-64. 
Wagner, D. H. 1993. Polystichum. Pages 290-299 in Flora 
of North America North of Mexico — Volume 3. Magno- 
liophyta: Magnoliidae and Hamamelidae. Edited by Flora 
of North America Editorial Committee. Oxford University 
Press Inc. New York, New York. 475 pages. 

Walker, T. G. 1979. The cytogenetics of ferns. In The experi- 
mental Biology of ferns. Edited by A. F. Dyer. Academic 
Press, London, England. 


Received 15 October 2002 
Accepted 1 September 2004 


Conservation Evaluation of the Pacific Population of Tall Woolly-heads, 
Psilocarphus elatior, an Endangered Herb in Canada* 


GEORGE W. DouGLas! and JEANNE M. ILLINGWORTH? 


‘Conservation Data Centre, British Columbia Ministry of Sustainable Resource Management, Terrestrial Information Branch, 
P.O. Box 9993 Stn Prov Govt, Victoria, British Columbia V8W 9R7 Canada 

23537 Savannah Ave., Victoria, British Columbia V8X 1S6 Canada 

Current address: Douglas Ecological Consultants Ltd., 6230 North Road, Duncan, British Columbia V9L 6K9 Canada 


Douglas, George W., and Jeanne M. Illingworth. 2004. Conservation evaluation of the Pacific population of Tall Woolly- 
heads, Psilocarphus elatior, an endangered herb in Canada. Canadian Field-Naturalist 118(2): 169-173. 


In Canada, Psilocarphus elatior occurs in British Columbia, Alberta and Saskatchewan. This paper examines the status of 
the Pacific populations located on southeastern Vancouver Island in southwestern British Columbia. The Pacific population 
consists of 12 recorded sites of which only five have been confirmed since 1993. In British Columbia, P. elatior is 
associated with dried beds of vernal pools and other open, moist depressions at lower elevations. In British Columbia, P. 


elatior populations occur in large numbers at only two of the seven locations. 


Key Words: Tall Woolly-heads, Psilocarphus elatior, endangered, distribution, population size, British Columbia. 


Tall Woolly-heads, Psilocarphus elatior (A. Gray) 
A. Gray” [Taxonomy and nomenclature follows Doug- 
las (1998) and Douglas et al. (1998; 1999a, c; 2001a 
b), is a member of a genus of five species which occur 
in the Americas (Cronquist 1950). It is one of three 
species occurring in British Columbia and Canada 
(Douglas 1998). 

Psilocarphus elatior is a small, erect, annual plant 
up to 15 cm tall (Figure 1; Douglas 1995, 1998). The 
opposite leaves are silky-hairy, linear-oblong, entire 
on the margins and 1.2 to 3.5 mm long. The flowers 
form solitary spherical heads in the leaf axils or at the 
tips of the stems or leaf branches. The flowers lack 
involucres but have involucral-like leaves at their bases. 
Each of the outer (female) threadlike ray flowers has 
a well developed, 2.4 to 3.8 mm long receptacle bract. 
These bracts have a translucent appendage below the 
summit. 

In British Columbia, P. elatior may be distinguished 
from Slender Woolly-heads (P. tenellus var. tenellus) 
by its erect habit, larger heads and receptacular bracts. 
In the field, young specimens of P. elatior may also be 
confused with young specimens of Lowland Cudweed 
(Gnaphalium palustre). Close examination will reveal 


* Field work for the Psilocarphus elatior project was fund- 
ed by the British Columbia Conservation Data Centre. 
The results appear in the British Columbia Conservation 
Data Centre database and a rare plant manual (Douglas et 
al. 2002). These data formed the basis for a Committee on 
the Status of Endangered Wildlife in Canada status report 
(Douglas, et al. 1999b) and the subsequent assessment of 
endangered (COSEWIC 2004*) for the Pacific population. 
The present paper also includes more recent information 
from research funded by the British Columbia Conserva- 
tion Data Centre. 


that most of the leaves of the latter are alternate and the 
flower heads have densely woolly involucres, broader 
ray flowers and a non-spherical shape. 


North American and Provincial Ranges 
Psilocarphus elatior ranges from southwestern Brit- 
ish Columbia, southeastern Alberta and southwestern 
Saskatchewan in Canada, south through Idaho, Wash- 
ington and Oregon to northern California (Cronquist 
1955; Scoggan 1979; Morefield 1993; Douglas 1998). 
In British Columbia, it is restricted to south-eastern 
Vancouver Island in south-western British Columbia 
(Figure 2; Douglas 1998; Douglas et al. 2002). 


Habitat 

Psilocarphus elatior inhabits dried beds of vernal 
pools, ephemeral lake edges (Figure 3) and other open 
moist, often disturbed sites at lower elevations. The 
sites are often level and generally slightly depressed. 
Psilocarphus elatior is not found within particular 
communities and often occupies sites where other spe- 
cies are sparse. There does not appear to be a consis- 
tent association with other plants. 

The Somenos Lake locality near Duncan supports 
the largest and most vigourous population of P. elatior 
(Table 1). The plants grow on the moist shoreline of 
the lake and appear when the lake level recedes in late 
spring or early summer. Major associates at this site 
include Slender-beaked Sedge (Carex athrostachya), 
One-sided Sedge (C. unilateralis), Brass Buttons 
(Cotula coronopifolia) and Skunkweed (Navarretia 
squarrosa). 

At Uplands Park, in the Municipality of Uplands 
near Victoria, five subpopulations occur in a large 
ephemeral meadow. Associated species include Bent- 
grass (Agrostis sp.), California Oatgrass (Danthonia 


169 


170 THE CANADIAN FIELD-NATURALIST 


Figure 1. Illustration of Psilocarphus elatior (line drawing 
by Elizabeth J. Stephen in Douglas [1995, 1998]). 


californica), Toad Rush (Juncus bufonis), Perennial 
Ryegrass (Lolium perenne) and Small Hop-clover 
(Trifolium dubium). Introduced species are more abun- 
dant here than at the Somenos Lake site and appear 
to be increasing yearly. 

The Christmas Hill site, although smaller than the 
previous two sites, is in relatively good condition. 
Native species in this habitat include Carolina 
Meadow-foxtail (Alopecurus carolinianus), Green- 
sheathed Sedge (Carex feta), Heterocodon (Hetero- 
codon rariflorum), and Scouler’s Popcornflower 
(Plagiobothrys scouleri). 

The two remaining populations, at Scafe Hill and 
Cattle Point, occur in small depressions. These popu- 
lations are probably the least stable of the extant pop- 
ulations. The Cattle Point populations are absent dur- 
ing some years but the seed bank appears to remain. 

Psilocarphus elatior is considered to be a vernal 
pool specialist (Keeley and Zedler 1998). They define 
vernal pools as “precipitation-filled seasonal wetlands 
inundated during periods when temperature is suffi- 
cient for plant growth, followed by a brief water- 
logged-terrestrial stage and culminating in extreme 
desiccating soil conditions of extended duration”. The 


Vol. 118 


f ss 
L PORT 

VANCOUVER “iw ‘fp 
ISLAND 


O 
UCLUELET 


SIDNEY 


FIGURE 2. The location and status of Psilocarphus elatior 
sites in British Columbia (0 — extirpated sites, e — re- 
cently confirmed sites, Hi — present status unknown). 


species is able to outcompete grassland species due 
to its tolerance of inundation and aquatic/wetland 
species due to its tolerance of soil desiccation and heat 
during summer drought. 


Biology 

Other than general habitat information, there is little 
in the literature regarding the biology and ecology of 
Psilocarphus elatior. It is likely, however, that this 
plant shares many of the same traits that are typical 
of the genus. 


2004 


DOUGLAS AND ILLINGWORTH: PACIFIC POPULATION OF TALL WOOLLY-HEADS 17] 


Ficure 3. A dense population of Psilocarphus elatior on the edge of Somenos Lake. The prominent sedge is Carex athrostachya. 
This site is usually submerged until late spring. 


Lack of structures attractive to insects and animals, 
and an interpretation of the floral structure indicate 
these inconspicuous, woolly annuals may self-polli- 
nate. Cronquist (1950) suggested that in the genus 
Psilocarphus, the position of the receptacular bracts, 
together with the position of the corolla and stigmas, 
effectively guide these latter structures towards the 
central flowers. Since it is the central flowers alone that 
produce pollen, and since the wool and leaves sur- 


rounding the head appeared to prevent pollen loss by 
wind, self-pollination is indicated. It is also possible 
that pollen may not be essential for seed-production 
and that asexual reproduction may be occurring, al- 
though chromosome counts would be necessary to 
verify this. 

Seed dispersal also appears limited. Since the ach- 
enes are much smaller than the enclosing bracts, the 
only obvious means of dispersal is by water or wind. 


Table 1. Locations and Population Sizes for Psilocarphus elatior in British Columbia. 


Collection Site Last 
Observation 
Cloverdale (Victoria) 1887 
Cedar Hill (Victoria) 1887 
Ucluelet 1909 
Roberts Bay (Sidney) 1913 
Swartz Bay (Sidney) 1931 
Francis-King Regional Park (Victoria) 1962 
University of Victoria (Victoria) 1966 
Cattle Point, Uplands Park (Victoria) 1993 
Scafe Hill (Victoria) 1996 
Uplands Park (Victoria) 1998 
Christmas Hill (Victoria) 2001 
Somenos Lake (Duncan) 2002 


Collector/ Population 
Observer (no./area) 
Macoun Extirpated 
Macoun Extirpated 
Macoun Extirpated 
Macoun Extirpated 
Groh Extirpated 
Melburn Unknown 
Turner Extirpated 
Ryan 200/10 m? 
Roemer 20/? m? 
Douglas 40 000+/1 200+ m? 
Douglas & Penny 450/72 m? 
Douglas & Douglas 100 000+/45 m? 


172 


The plant’s habitat in vernal pools may also permit the 
bracts and achenes to be transferred by the muddy 
feet of waterfowl and other animals (Cronquist 1950). 

Further studies are required to determine many as- 
pects of the population dynamics of P. elatior includ- 
ing the average life-cycle of the species, the frequency 
and requirements for seed germination and survival, 
and its competitive ability with other species. 


Population Attributes 

Psilocarphus elatior has been recorded from 12 sites 
in southwestern British Columbia (Table 1). Five of 
these have been confirmed since 1993. The status of 
the remaining populations is unknown and many are 
believed extirpated. Population areas range from 10 m? 
to over 1200 m? while numbers of plants range from 
20 to over 100 000. In the majority of cases, popula- 
tion trend analysis is not available because of limited 
demographic data. 


Provincial, National and Global Ranks 

Provincially, P. elatior is ranked S2 by the British 
Columbia Conservation Data Centre (Douglas et al. 
2002) which indicates this species to be “imperiled 
because of rarity (typically 6-20 extant occurrences or 
few remaining individuals) or because of some fac- 
tor(s) making it vulnerable to extirpation or extinction.” 
Nationally the species is ranked N3, while globally 
it has a rank of G5. The latter rank indicates it is 
“frequent to common to very common; demonstrably 
secure and essentially ineradicable under present con- 
ditions”. 


Threats and Protection 

Habitat destruction is the greatest threat to the exist- 
ing populations of Psilocarphus elatior. Some sites 
occur in areas subjected to heavy pedestrian tram- 
pling or are at risk from private development. Threats 
to the continued survival of this species are compound- 
ed by the lack of biological and ecological information 
which create difficulties in site management. 

Four of the P. elatior populations are partially pro- 
tected by their location in municipal or regional parks 
or special protected areas (Somenos Lake). The latter 
probably has the greatest degree of protection since it 
is administered by British Columbia Parks and falls 
under the Park Act. Most of the municipal or regional 
parks receive little active management at the present 
time, at least with respect to their rare plants. Park 
enhancement projects, road and trail developments and 
heavy recreational use by humans often result in the 
destruction of the native vegetation and rare plant spe- 
cies. Psilocarphus elatior could be a candidate species 
for protection under the provincial Wildlife Amendment 
Act as it is currently Red-listed by the British Colum- 
bia Conservation Data Centre (Douglas et al. 2002). 

Most of the populations contain small numbers of 
plants. Once a population becomes small, it becomes 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


more susceptible to demographic and environmental 
variation and loss of genetic variability. In some cases, 
small populations are at risk of inbreeding depres- 
sion, genetic drift and loss of fitness (Primack 1998). 


Evaluation 

The British Columbia Conservation Data Centre 
considers Psilocarphus elatior to be threatened/endan- 
gered in British Columbia (Douglas et al. 2002a) and 
the Committee on the Status of Endangered Wildlife 
in Canada has assessed the species as endangered 
(COSEWIC 2003). Most of the populations are small 
and not viable. With limited knowledge of the plants 
biological and ecological requirements, this species is 
vulnerable to extirpation in British Columbia. Without 
research on growth requirements and further demo- 
graphic information, the stability of the present popu- 
lations will remain unknown. Studies are also neces- 
sary to determine if the present habitats are necessary 
for the successful growth of P. elatior or if these plants 
are simply outcompeted elsewhere. The limited num- 
ber of individuals also reduces the potential for gen- 
etic variation which may be necessary to respond to 
environmental changes in the future. 


Acknowledgments 
We thank Sylvia Douglas, Jenifer Penny, and 
Michael Ryan for their field assistance. 


Documents Cited (marked * in text) 

COSEWIC. 2004. Canadian Species at Risk. Committee on 
the Status of Endangered Wildlife in Canada. 
www.cose wic.gc.ca, Ottawa, Ontario. 13 pages. 

Douglas, G. W., J. Gould, and J. M. Illingworth. 1999. Status 
report on the Tall woolly-heads, Psilocarphus elatior (A. 
Gray) A. Gray. Unpublished report. Committee on the 
Status of Endangered Wildlife in Canada, Ottawa. 19 pages. 


Literature Cited 

Cronquist, A. 1950. A review of the genus Psilocarphus. 
Research Studies of the State College of Washington 18: 
71-89. 

Cronquist, A. 1955. Vascular plants of the Pacific Northwest. 
Part 5: Compositae. University of Washington Press, Seat- 
tle, Washington. 343 pages. 

Douglas, G. W. 1995. The sunflower family (Asteraceae) of 
British Columbia. Volume 2 — Astereae, Anthemideae, 
Eupatoreae and Inuleae. Royal British Columbia Museum, 
Victoria, British Columbia. 393 pages. 

Douglas, G. W. 1998. Asteraceae. Pages 96-392 in Illustrat- 
ed flora of British Columbia. Volume 1. Gymnosperms and 
Dicotyledons (Aceraceae through Asteraceae). Edited by 
G. W. Douglas, G. B. Straley, D. Meidinger, and J. Pojar. 
British Columbia Ministry of Environment, Lands and 
Parks and British Columbia Ministry of Forests, Victoria, 
British Columbia. 

Douglas, G. W., D. Meidinger, and J. L. Penny. 2002. Rare 
native vascular plants of British Columbia. Second edition. 
Province of British Columbia, Victoria, British Columbia. 
358 pages. 


2004 


Douglas, G. W., D. Meidinger, and J. Pojar. 1999a. Illus- 
trated flora of British Columbia. Volume 3. Dicotyledons 
(Diapensiaceae through Onagraceae). British Columbia 
Ministry of Environment, Lands and Parks and British 
Columbia Ministry of Forests, Victoria, British Columbia. 

Douglas, G. W., D. Meidinger, and J. Pojar. 1999b. Illus- 
trated flora of British Columbia. Volume 4. Dicotyledons 
(Orobanchaceae through Rubiaceae). British Columbia 
Ministry of Environment, Lands and Parks and British Col- 
umbia Ministry of Forests, Victoria, British Columbia. 
427 pages. 

Douglas, G. W., D. Meidinger, and J. Pojar 2001a. Illus- 
trated flora of British Columbia. Volume 6. Monocotyle- 
dons (Acoraceae through Najadaceae). British Columbia 
Ministry of Sustainable Resource Management and British 
Columbia Ministry of Forests, Victoria, British Colum- 
bia. 361 pages. 

Douglas, G. W., D. Meidinger, and J. Pojar 2001b. Ilus- 
trated flora of British Columbia. Volume 7. Monocotyle- 
dons (Orchidaceae to Zosteraceae). British Columbia 
Ministry of Sustainable Resource Management and British 
Columbia Ministry of Forests, Victoria, British Columbia. 
379 pages. 

Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar. 
1998. Illustrated flora of British Columbia. Volume 1. 


DOUGLAS AND ILLINGWORTH: PACIFIC POPULATION OF TALL WOOLLY-HEADS 


173 


Gymnosperms and Dicotyledons (Aceraceae through 
Asteraceae). British Columbia Ministry of Environment, 
Lands and Parks and British Columbia Ministry of Forests, 
Victoria, British Columbia. 436 pages. 

Keeley, J. E., and P. H. Zedler. 1998. Characterization and 
global distribution of vernal pools. /n Ecology, conserva- 
tion, and management of vernal pool ecosystems: Proceed- 
ings from a 1996 Conference. Edited by C. W. Witham, 
E. T. Bauder, D. Belk, W. R. Ferren Jr., and R. Ornduff. 
California Native Plant Society. Sacramento, California. 

Morefield, J. 1993. Psilocarphus. Page 329 in The Jepson 
manual: higher plants of California. Edited by J. C. Hick- 
man. University of California Press, Berkeley, California. 
1400 pages. 

Primack, R. B. 1998. Essentials of Conservation Biology. 
24 Edition. Sinauer Associates Inc. Sunderland, Massa- 
chusetts. 

Scoggan, H. J. 1979. The flora of Canada. Part 4 — Dico- 
tyledoneae (Loasaceae to Compositae. National Museum 
of Natural Sciences, Ottawa, Ontario. Publication in Botany 
Number 7. Pages 1117-1711. 


Received 15 October 2002 
Accepted | September 2004 


Conservation Evaluation of Howell’s Triteleia, Triteleia howellii, an 
Endangered Lily in Canada* 


GEORGE W. DouGLas!:? and JENIFER L. PENNY! 


‘Conservation Data Centre, British Columbia Ministry of Sustainable Resource Management, Terrestrial Information 
Branch, P.O. Box 9993 Stn Prov Govt, Victoria British Columbia V8W 9R7 Canada 
? Current address: Douglas Ecological Consultants Ltd., 6230 North Road, Duncan, British Columbia V9L 6K9 Canada 


Douglas, George W., and Jenifer L. Penny. 2004. Conservation evaluation of Howell’s Triteleia, Triteleia howellii, an 
endangered lily in Canada. Canadian Field-Naturalist 118(2): 174-178. 


In Canada, Triteleia howellii is restricted to Quercus garryana stands and grass-dominated meadows on southeastern Van- 
couver Island in southwestern British Columbia. Nine sites have been confirmed in recent years while three other sites are 
considered extirpated. These Canadian sites represent the northern range limits of 7: howellii. Threats to existing populations 
vary in intensity. Although most populations are protected to a certain extent from direct habitat destruction, introduced species 
pose a serious potential threat to the continued existence of most populations. Managing sites for 7: howellii is difficult because 


little information is available regarding the general biology of this species. 


Key Words: Howell’s Triteleia, Triteleia howellii, endangered, distribution, population size, British Columbia. 


Howell’s Triteleia, Triteleia howellii (S. Wats.) 
Greene [taxonomy and nomenclature follow Douglas 
et al. (1998a, b; 1999a, b; 2000, 2001a, b)], some- 
times treated as T. grandiflora Lindl. var. howellii (S. 
Wats.) Hoover, is a member of a genus of 14 species 
in North America (Keator 1993). Three species occur 
in British Columbia and Canada (Scoggan 1979; Pojar 
2001). 

Triteleia howellii is a perennial herb from a deep, 
straw-coloured, fibrous-scaly, nearly globe-shaped, 
bulb-like corm (Figure 1; Pojar 2001). The erect, flow- 
ering stem is 20-50 cm tall with one or two smooth, 
slender, linear basal leaves. The leaves are 20-40 cm 
long, 3-8 mm wide, sheathed at the base and have en- 
tire margins. The flowers consist of six whitish to blue, 
vase-shaped to narrowly bell-shaped, fused segments 
forming a 1.5-2 cm long tube. The corolla lobes, which 
are about as long as the tube, are in two, spreading, 
petal-like whorls, about as long as the tube. The outer 
three are broadly lanceolate, the inner three are oblong- 
egg-shaped and all are slightly ruffled. The fruit con- 
sists of a stalked, egg-shaped capsule containing black 
rounded seeds. 

Triteleia howellii is similar in appearance to its close 
relative 7: grandiflora. It is distinguished from the latter 
* The field work for the Triteleia howellii project was funded 

by the British Columbia Conservation Data Centre. The 

results appear in the British Columbia Conservation Data 

Centre database and a rare plant manual (Douglas et al. 

2002). This information formed the basis for a Committee 

on the Status of Endangered Wildlife in Canada status 

report (Douglas 2003*) and the subsequent assessment of 

Endangered (COSEWIC 2003*). The present paper also 

includes more recent information, funded by the Nature 

Conservancy of Canada and the Habitat Conservation Trust 

Fund, that will be used in a National Recovery Strategy 

for T: howellii (Douglas and Smith 2005). 


by its flat filaments which are attached at the same level 
on the perianth tube (Pojar 2001). The filaments of 7: 
grandiflora, in contrast, are not flat and are attached 
at two levels on the perianth tube. 


North American and Provincial Ranges 

Triteleia howellii ranges from southwestern British 
Columbia, south through Washington and Oregon to 
northern California (Barkworth 1977a; Keator 1993). 
In Canada, 7: howellii is known only from southeast- 
ern Vancouver Island (Figure 2; Pojar 2001; Douglas 
et al. 2002). 


Habitat 

In British Columbia T- howellii occurs on rock out- 
crops, in Garry Oak (Quercus garryana) woodlands, 
Garry Oak/Arbutus (Arbutus menziesii) stands and 
occasionally in highly disturbed sites dominated by 
weeds in private yards and on roadsides. In the highly 
disturbed sites, dominants include Orchard Grass (Dac- 
tylis glomerata), Cheat Grass (Bromus tectorum), Com- 
mon Vetch (Vicia sativa), Rip-gut Brome (Bromus 
rigidus), Soft Brome (B. hordeaceus), Perennial Rye- 
grass (Lolium perenne), and Pacific Sanicle (Sanicula 
crassicaulis var. crassicaulis), all introduced except 
for Sanicula. 

At the highest quality site, in the Quercus garryana 
woodland at the Cowichan Garry Oak Preserve, the 
habitat is classified as a Quercus garryana/Dactylis 
glomerata plant community (Douglas et al. 2002*) 
and is characterized by deep, dark soils up to a metre in 
depth. It is likely that prior to understory dominance 
by D. glomerata in this (Figure 3), and other Quercus 
stands of the region, this plant community would have 
fallen within the Q. garryana/California Brome (Bro- 
mus carinatus) community type (Roemer 1972). An 


174 


2004 


FiGure 1. Illustration of Triteleia howellii (Line drawing in 
Pojar 2001). 


extremely rich low shrub and herb stratum is present 
during the spring. The most prominent species in the 
Cowichan Garry Oak Preserve Garry Oak stand are 
Sanicula crassicaulis var. crassicaulis and Dactylis 
glomerata (Douglas et al. 2002*). Other species with 


DOUGLAS AND PENNY: HOWELL’S TRITELEIA IN CANADA 


Wes 


x 


Ay 9 
Fie: \ comoxS SER 3 
ie Nil ¢ 


AV f =, South 
g T Wellington 
NANAIMO*\4> 
\ 


VANCOUVER 
ISLAND 


FiGureE 2. Distribution of Triteleia howellii in British Col- 
umbia (o — extirpated sites, e — recently confirmed 
sites). 


moderate to high constancies associated with T. how- 
ellii include Common Camas (Camassia quamash), 
Bromus spp., Broad-leaved Shooting-star (Dodecatheon 
hendersonii_ ssp. hendersonii), Cleavers (Galium 
aparine), and Common Snowberry (Symphoricarpos 
albus). A marked change in composition takes place 
by mid-summer. Many of the conspicuous native plants 
(e.g., Great Camas (Camassia leichtlinii), C. quamash, 
Dodecatheon hendersonii, and Yellow Montane Violet 
(Viola praemorsa ssp. praemorsa)) have completed 
their yearly life cycle and have essentially disappeared. 
Perennial grasses that were not recognizable or had 
not initiated growth in the spring and numerous intro- 


176 


THE CANADIAN FIELD-NATURALIST 


TABLE |. Locations and population sizes for Triteleia howellii in Canada 


Collection Last 
Site Observation 
Oak Bay (Victoria) 1912 
Saanich Arm (Victoria) 1919 
Witty’s Lagoon Regional Park (Metchosin) 1999 
Gordon Head (Saanich) 1999 
Cowichan Garry Oak Preserve (Duncan) 1999 
Cowichan River Estuary (Duncan) 2001 
Thetis Lake Regional Park (View Royal) 2002 
Uplands Park (Victoria) 2003 
Mt. Tzuhalem, base of (along Khenipsen Road) 2003 
Albert Head Lagoon Regional Park (Metchosin) 2003 
Horth Hill Regional Park (North Saanich) 2003 
Beacon Hill Park (Victoria) 2004 
Somenos Lake (Duncan) 2004 
Williams Head Road (Metchosin) 2004 


duced annuals, well adapted to the drier soils, domi- 
nate the understory. At this time, Dactylis glomerata 
and Vicia species are the most prominent species 
with greatly increased mean covers. Other prominent 
species in mid-summer include the native grasses, 
Bromus carinatus and Alaska Oniongrass (Melica 
subulata), and the introduced grasses, Barren Brome 
(Bromus sterilis) and Kentucky Bluegrass (Poa pra- 
tensis). 

Triteleia howellii also occurs in a Quercus garryana 
— Arbutus menziesii stand. The shrub layer is more 
prominent at this site and is dominated by Tall Oregon- 
grape (Mahonia aquifolium) and Oceanspray (Holo- 
discus discolor). Major associates include Hairy 
Honeysuckle (Lonicera hispidula), Bromus rigidus, 
Galium aparine, Small-flowered Nemophila (Nemo- 
phila parviflora) and Hedgehog Dogtail (Cynosurus 
echinatus). 


Biology 

There is little information known about the biology 
or ecology of Triteleia howellii throughout its range. 
Reproduction is by division of the corm, by produc- 
tion of numerous cormlets, and by seed (Barkworth 
1977b). 


Population Attributes 

Triteleia howellii has been collected at 14 sites in 
Canada, all of which are located on southeastern 
Vancouver Island (Table 1). Of the 14 sites, 11 have 
been confirmed since 1999 while the status of the re- 
maining three sites is unknown and are likely extir- 
pated. Population areas range from small (one m7) to 
over three or four hectares, while plant numbers range 
from a single plant to over 430 plants (Table 1). Little 
information is available on population trends. The sites 
that have recently been examined show the populations 


Vol. 118 
Collector/ Population 
Observer (no./area) 

Beaven Extirpated 

Newcombe Extirpated 

Douglas & Penny 43/200 m? 

Fontaine 51/5 m? 

Douglas 430/3-4 ha 

Douglas 62/3 m? 

Ceska 1/1 m?* 

Penny & Fairbarns Not seen since 1917, 
2003 search 
unsuccessful, probably 
extirpated 

Janszen 6/.05 m?* 

Roemer 8/2 m? 

Janszen 3/1 m?* 

Fairbarns ca 200/12 m? 

Douglas & Richards 90/140 m? 

Milne 14/?m? 


are apparently stable although numbers of flowering 
plants may vary. 


Provincial, National and Global Ranks 

The British Columbia Conservation Data Centre has 
ranked this species S2 and placed it on the British 
Columbia Ministry of Sustainable Resource Manage- 
ment red-list (Douglas et al. 2002). This is the most 
critical category for imperilled rare native vascular 
plants in British Columbia. A rank of S2 is consid- 
ered “critically imperilled because of rarity (typically 
6-20 extant occurrences or few remaining individuals) 
or because of some factor(s) making it vulnerable to 
extirpation or extinction.” Since the species is restrict- 
ed to British Columbia, the National rank is N2. Glo- 
bally, Triteleia howellii is ranked G3G4, indicating 
that, although the presently known sites are less than 
100, it is more likely that this species is frequent to 
common in its range and apparently secure. 


Threats and Protection 

The most direct and immediate threat to Triteleia 
howellii is habitat destruction. This is of particular con- 
cern in the grass-dominated meadows often associ- 
ated with the Quercus garryana communities that are 
restricted to the southeastern side of Vancouver Island 
and some of the Gulf Islands. This type of vegetation 
was much more common before colonization by 
European settlers. This destruction has continued to 
the present resulting in the elimination of almost all 
sites occurring outside parks or ecological reserves. 
Historically, Q. garryana communities and grass- 
dominated meadows have always been heavily influ- 
enced by human activity, especially fires. Roemer 
(1972) believed that without human interference some 
of these stands would have eventually been replaced 
by Douglas-fir forests. 


2004 


DOUGLAS AND PENNY: HOWELL’S TRITELEIA IN CANADA 


177 


FiGure 3. Habitat of Triteleia howellii in a Quercus garryana stand in the Cowichan Garry Oak Preserve near Quamichan 
Lake. Dactylis glomerata is the dominant grass in this late summer photo. 


The suppression of fire within the past century may 
also have contributed to the decrease of Triteleia how- 
ellii populations. Most of the sites in which T: howellii 
has been collected were likely maintained in the past 
as a result of periodic fires, both natural and unnatural. 
In the past, aboriginal peoples probably set fire to 
these stands to maintain them as an important habitat 
for wildlife (Roemer 1972). Since that time, these sites 
have experienced little disturbance, resulting in the in- 
vasion and expansion of many other species, especial- 
ly introducted grasses. 

The introduction of European species has resulted 
in substantial, and probably irreversible, changes not 
only to the grass-dominated meadows associated with 
Quercus garryana, but also to the rocky xeric sites 
north and west of Victoria where Triteleia howellii 
has been collected in the past. One of the most devas- 
tating species is Scotch Broom (Cytisus scoparius), 
which has become a dominant shrub on xeric, exposed 
sites throughout much of southeastern Vancouver 
Island and the Gulf Islands. Much of the vegetation 
is now dominated by introduced grasses. These species 
include Early Hairgrass (Aira praecox), Sweet Vernal- 
grass (Anthoxanthum odoratum), Cynosurus echinatus 
and Dactylis glomerata. 

Some of the populations contain very few plants. 
Once a population becomes small, it becomes more 


vulnerable to demographic and environmental vari- 
ation and loss of genetic variability. In some cases, 
small populations are at risk of inbreeding depression, 
genetic drift and loss of fitness (Primack 1998). 

The population with the best protection is located at 
the Cowichan Garry Oak Preserve where the general 
public is excluded. This five-hectare stand near Quami- 
chan Lake on Vancouver Island represents the best 
example of a Quercus garryana woodland in the prov- 
ince and probably one of the best in the Pacific North- 
west. The stand contains a relatively low number of 
exotic species and has active management plans in 
place. This site also contains the largest known popu- 
lation of T; howellii in the province (Douglas et al. 
2002*). 

A number of Triteleia howellii populations are also 
in small regional parks in the Greater Victoria area. 
These include populations at Beacon Hill, Witty’s 
Lagoon, Thetis Lake, and Horth Hill. Most of these 
parks receive little active management at the present 
time, at least with respect to their rare plants. Park 
enhancement projects, road and trail developments 
and heavy recreational use by humans often result in 
the destruction of the native vegetation and rare plant 
species. 

Triteleia howellii could be a candidate species for 
protection under the provincial Wildlife Amendment 


178 


Act as it is currently red-listed by the British Colum- 
bia Conservation Data Centre. One of the populations 
(Somenos Lake) of T: howellii is protected by the Park 
Act since it is currently administered by Parks BC. 


Evaluation 

The British Columbia Conservation Data Centre 
considers Triteleia howellii to be threatened/endangered 
in British Columbia (Douglas et al. 2002a) and the 
Committee on the Status of Endangered Wildlife in 
Canada has assessed the species as endangered 
(COSEWIC 2003). About 1000 flowering plants have 
been recorded in recent years from 11 locations. Only 
four of these populations can be considered viable since 
the remaining populations cover areas of less than 5 m?. 
The prognosis for this species may not be good since, 
since only one population, at the Cowichan Garry Oak 
Preserve, is included in an active management plan. 


Acknowledgments 

We thank Sharon Hartwell, Marie Fontaine, and 
Lora May Richards for their assistance with field work. 
We also thank Adolf and Oluna Ceska, Matt Fairbarns, 
Moralea Milne and Hans Roemer for information on 
populations at several sites. 


Documents Cited 

COSEWIC. 2003. COSEWIC assessment and status report 
on Howell’s Triteleia, Triteleia howellii in Canada. Com- 
mittee on the Status of Endangered Wildlife in Canada. 
www.cosewic.gc.ca, Ottawa, Ontario. 16 pages 

Douglas, G. W. 2003. COSEWIC Status report on Howell’s 
Triteleia, Triteleia howellii in Canada, in COSEWIC as- 
sessment and status report on Howell’s Triteleia, Triteleia 
howellii in Canada. Committee on the Status of Endan- 
gered Wildlife in Canada. www.cosewic.gc.ca, Ottawa, 
Ontario. 16 pages. 

Douglas, G. W., J. L. Penny, and R. E. Maxwell. 2002b. 
Composition, phenology, stand structure and soils of a 
Quercus garryana (Garry Oak) woodland community at 
Quamichan Lake, Vancouver Island, British Columbia. 
British Columbia Conservation Data Centre, Ministry of 
Sustainable Resource Management, Victoria, British 
Columbia. 52 pages. 

Douglas, G. W., and S. J. Smith. 2005. National multi-species 
recovery strategy for species at risk in Garry Oak wood- 
lands. In preparation. Ottawa, Ontario. 


Literature Cited 

Barkworth, M. E. 1977a. The taxonomic status of Brodi- 
aea howellii Watson (Liliaceae). Northwest Science 51: 
139-148. 

Barkworth, M. E. 1977b. Intraspecific variation in Brodi- 
aea douglasii Watson (Liliaceae). Northwest Science 51: 
79-90. 

Douglas, G. W., D. Meidinger, and J. L. Penny. 2002a. 
Rare native vascular plants of British Columbia. Second 
edition. Province of British Columbia. Victoria, British 
Columbia. 359 pages. 

Douglas, G. W., D. Meidinger, and J. Pojar. 1999a. Illus- 
trated flora of British Columbia. Volume 3. Dicotyledons 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


(Diapensiaceae through Onagraceae). British Columbia 
Ministry of Environment, Lands and Parks and British 
Columbia Ministry of Forests, Victoria, British Colum- 
bia. 423 pages. 

Douglas, G. W., D. Meidinger, and J. Pojar. 1999b. Illus- 
trated flora of British Columbia. Volume 4. Dicotyledons 
(Orobanchaceae through Rubiaceae). British Columbia 
Ministry of Environment, Lands and Parks and British 
Columbia Ministry of Forests, Victoria, British Colum- 
bia. 427 pages. 

Douglas, G. W., D. Meidinger, and J. Pojar. 2000. Illus- 
trated flora of British Columbia. Volume 5. Dicotyledons 
(Salicaceae to Zygophyllaceae) and Pteridophytes. British 
Columbia Ministry of Environment, Lands and Parks and 
British Columbia Ministry of Forests, Victoria, British 
Columbia. 389 pages. 

Douglas, G. W., D. Meidinger, and J. Pojar. 2001a. Illus- 
trated flora of British Columbia. Volume 6. Monoco- 
tyledons (Acoraceae to Najadaceae). British Columbia 
Ministry of Environment, Lands and Parks and British 
Columbia Ministry of Forests. Victoria, British Columbia. 
361 pages. 

Douglas, G. W., D. Meidinger, and J. Pojar. 2001b. Illus- 
trated flora of British Columbia. Volume 7. Monocotyle- 
dons (Orchidaceae to Zosteraceae). British Columbia 
Ministry of Sustainable Resource Management and British 
Columbia Ministry of Forests, Victoria, British Columbia. 
379 pages. 

Douglas, G. W., G. B. Straley, and D. Meidinger. 1998a. 
Illustrated flora of British Columbia. Volume 1. Gymno- 
sperms and Dicotyledons. (Aceraceae through Astera- 
ceae). British Columbia Ministry of Environment, Lands 
and Parks and British Columbia Ministry of Forests, 
Victoria, British Columbia. 436 pages. 

Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar. 
1998b. Illustrated flora of 

British Columbia. Volume 2. Dicotyledons. (Balsaminaceae 
through Cuscutaceae). British Columbia Ministry of Envi- 
ronment, Lands and Parks and British Columbia Ministry 
of Forests, Victoria, British Columbia. 401 pages. 

Keator, G. 1993. Triteleia. Pages 1206-1208. In The Jepson 
manual: Higher plants of California. Edited by J. C. 
Hickman. University of California Press. Berkeley, Cali- 
fornia. 1400 pages. 

Pojar, J. 2001. Liliaceae. Pages 270-318. Jn Illustrated flora 
of British Columbia. Volume 6. Monocotyledons (Acora- 
ceae to Najadaceae). Edited by G. W. Douglas, D. Meid- 
inger, and J. Pojar. British Columbia Ministry of Environ- 
ment, Lands and Parks and British Columbia Ministry of 
Forests, Victoria, British Columbia. 361 pages. 

Primack, R. B. 1998. Essentials of Conservation Biology. 
2™4 Edition. Sinauer Associates Inc. Sunderland, Massa- 
chusetts. 

Roemer, H. L. 1972. Forest vegetation and environments of 
the Saanich Peninsula, Vancouver Island. Ph.D. thesis. Uni- 
versity of Victoria, Victoria, British Columbia. 405 pages. 

Scoggan, H. J. 1979. The flora of Canada. Part 4. Dicotyle- 
doneae (Loasaceae to Compositae). National Museum of 
Natural Sciences, Ottawa, Ontario. National Museum of 
Natural Sciences Publications in Botany (7): 1117-1710. 


Received 15 October 2002 
Accepted | September 2004 


Conservation Evaluation of Small-flowered Lipocarpha, Lipocarpha 
micrantha (Cyperaceae), in Canada 


TYLER W. SmITH!, GEORGE W. DOUGLAS? AND ALLAN G. HARRIS? 


'Royal Botanical Gardens, P. O. Box 399, Hamilton, Ontario L8N 3H8 Canada 

Present address: Plant Science, McGill University, Raymond Building, 21 111 Lakeshore Road, Ste. Anne de Bellevue, 
Quebec H9X 3V9 Canada 

Conservation Data Centre, British Columbia Ministry of Sustainable Resource Management, Terrestrial Information 
Branch, P. O. Box 9993 Station Provincial Government, Victoria, British Columbia, V8W 9R7 Canada 

Present address: Douglas Ecological Consultants Ltd., 62030 North Road, Duncan, British Columbia V9L 6K5 Canada 

3Northern Bioscience, 136 S. Hill Street, Thunder Bay, Ontario P7B 3V1 Canada 


Smith, Tyler W., George W. Douglas, and Allan G. Harris. 2004 Conservation Evaluation of Small-flowered Lipocarpha, 
Lipocarpha micrantha (Cyperaceae), in Canada. Canadian Field-Naturalist 118(2): 179-184. 


In Canada, Lipocarpha micrantha has been documented at eight locations in Quebec, Ontario, and British Columbia. Four of 
these populations have apparently been extirpated. The remaining populations, ranging from 120 to approximately 40000 
plants, are all northern disjuncts from the main range of this species. Threats to these populations include water level 
regulation and shoreline development. Considering the threats to the habitat of Lipocarpha micrantha, and the small size of 
most of the remaining populations, it has been designated an Endangered species in Canada. 


Key Words: Small-flowered Lipocarpha, Lipocarpha micrantha, British Columbia, Ontario, Quebec, endangered, distribution, 


population size. 


Small-flowered Lipocarpha, Lipocarpha micrantha 
(Vahl.) G. Tucker, is the only member of a mainly trop- 
ical genus to occur in Canada. While it appears under 
this name in the Flora of North America (Tucker 2002), 
some recent treatments place it in the genus Hemicar- 
pha Nees (Gleason and Cronquist 1991). Lipocarpha, 
including Hemicarpha, is generally accepted as a dis- 
tinct genus. However, taxonomic clarification of closely 
related genera in the Cyperaceae may require nomen- 
clatural revisions, pending the outcome of ongoing 
research (Muasya et al. 2002). 

Lipocarpha micrantha is a caespitose annual sedge, 
2-20 cm tall with narrow (0.5 mm) leaves up to 10 cm 
long. The 1 to 3 subsessile flower spikes are borne at 
the top of the stem. The spikes are ovoid, 2—6 mm long, 
with numerous perfect flowers concealed behind spi- 
rally imbricate scales. The inflorescence is subtended 
by 2 to 3 leaf-like bracts, the lowest of which looks 
like a continuation of the stem (Figure 1: Hitchcock 
et al. 1969; Gleason and Cronquist 1991; Douglas 
and Ceska 2001; Tucker 2002). 

The densely tufted plants resemble the seedlings of 
many other sedge species that occur in the same habitat. 
In Canada Lipocarpha micrantha is perhaps most simi- 
lar in aspect to Awned Cyperus (Cyperus squarrosus 
L.), another small annual sedge. Lipocarpha micrantha 
is distinguished from this and all other Cyperus spe- 
cies by its spirally arranged flowers. 


Biology 

Lipocarpha micrantha is only visible during a short 
period each year. It germinates in late summer, when 
dropping water levels expose the open sandy habitat 


it requires. Flowering and fruiting occur in August and 
September. High water levels may prevent germination 
from occurring, with the population remaining dormant 
until appropriate conditions occur. We do not know 
how long dormant seeds may remain viable, but field 
observations suggest they can persist at least two years. 
Migrating waterfowl may be a vector for long-distance 
dispersal. 


Distribution 

Lipocarpha micrantha ranges from Brazil to Canada. 
North of Mexico, the main range extends from Texas, 
north to northwestern Ontario, and east to the east coast 
of the United States. Disjunct populations have been 
recorded east and west of the main range, in Alabama, 
Arizona, British Columbia, California, Florida, Ken- 
tucky, Maine, New Mexico, New York, North Carolina, 
and Quebec (Tucker 2002). 

In Canada, Lipocarpha micrantha occurs at Osoyoos 
Lake and Okanagan Lake in British Columbia (Figure 
2), and at Rainy Lake and Lake of the Woods in north- 
western Ontario (Figure 3). Populations along the De- 
troit River in southern Ontario and along the north shore 
of Lake Champlain in Quebec have been extirpated. 


Habitat 

Lipocarpha micrantha grows on sandy beaches and 
interdunal swales that are subject to seasonal flooding, 
but are protected from high waves or strong currents. 
It is usually found in areas of very sparse vegetation, 
and apparently is intolerant of competition from other 
plant species. These habitat conditions are maintained 
by fluctuating water levels. While Lipocarpha micrantha 


179 


180 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Ficure 1. Illustration of Lipocarpha micrantha (Line drawing by permission from Hitchcock et al. 1969). 


requires seasonal low water levels to germinate and 
flower, periodic high water is required to prevent 
more vigorous species from dominating its shoreline 
habitat (see Keddy and Reznicek 1986 for a discussion 
of the relationship between water level fluctuations 
and wetland vegetation). 

Common associates of Lipocarpha micrantha in- 
clude a variety of Cyperus, Bidens, and Salix species. 
Cyperus squarrosus has been noted as an associate at 
every station in Canada. 


Population Trends and Limiting Factors 

Of eight documented populations (Ceska and Ceska 
1980; Oldham and Crins 1988; Sabourin et al. 1992*; 
Oldham 1996; Harris et al. 2000*; Oldham 2000; Smith 


et al. 2002*) of Lipocarpha micrantha in Canada, 
only four persist (Table 1). 

The Lake Champlain population, in Missisquoi Bay, 
Quebec, was discovered in 1953 (Louis-Alphonse 
3458, specimen at MT; herbarium acronyms follow 
Holmgren et al. 2003*), and was documented through- 
out the 1950s. However, a search in 1989 failed to find 
any plants (Sabourin et al. 1992*). Most recently, TWS 
searched the site in 2002 but did not find any plants 
despite the presence of appropriate habitat and moder- 
ate water levels. This site is adjacent to a campground 
and recreational activity may have contributed to the 
demise of this population. Water quality may also have 
been a factor, as a river flowing into Lake Champlain 
at this location smells strongly of raw sewage. 


2004 


—~ 
—— 
———> \ 


SMITH, DOUGLAS, AND HARRIS: SMALL-FLOWERED LIPOCARPHA 181 


KAMLOOPS 


SPENCES BRIDGE 


I 
i) aw 
m4 
7 


_) \ENDERBY 
_/ ARMSTRONG 


/ 
. Okanagan ; 


Z 
Te 


camoce 


KEREMEOS, 


BRITISH 
COLUMBIA 
NE 


Lake . j 


Ree 


NCETON ~y PENTICTON J 


On 


Ficure 2. Distribution (stars) of Lipocarpha micrantha in British Columbia. 


The oldest records of Lipocarpha micrantha in Can- 
ada are collections from the Detroit River shoreline, 
south of Windsor, Ontario, in 1892 (Macoun 28668, 
specimen at CAN) and 1901 (Macoun 7594, speci- 
mens at TRTE, GH). The populations represented by 
these collections have presumably been destroyed by 
shoreline development. The only recent records from 
this region are from the vicinity of Holiday Beach Con- 


servation Area, near Amherstburg. This population was 
discovered in 1984, when 15 plants were observed 
(Oldham and Crins 1988). The population was still 
present in 1987 (Sabourin et al 1992*), but no plants 
were found during two searches by TWS in 2001. The 
habitat at Holiday Beach has been seriously degraded: 
dense mats of algae covered the beach, except in areas 
where it had been cleared away by heavy machinery. 


182 THE CANADIAN FIELD-NATURALIST Vol. 118 
TABLE |. Locations and Population Sizes for Lipocarpha micrantha in Canada. 
Site Last Observer (Last Population 

Observation Searcher) (number/area) 

Osoyoos Lake (British Columbia) 1980 Ceska Extirpated 
Osoyoos Lake, 2001 Douglas 30 000-50 000/2.3ha 

Osoyoos Indian Reserve (British Columbia) 
Okanagan Lake, 2002 Klinkenberg 120/2 m?* 

Sun Oka Beach Provincial Park (British Columbia) 
Sable Island (Ontario) 2001 Harris 1 800/0.4ha 
Poundnet Bay (Ontario) 2000 (see text) Harris 75/0.00lha 
Holiday Beach (Ontario) 1987 (2001 ) Oldham (Smith) Extirpated 
Detroit River (Ontario) 1901 Macoun Extirpated 
Missisquoi Bay (Quebec) 1957 (2002 ) Louis-Alphonse Extirpated 

(Smith) 


The Lake of the Woods population, at Sable Island, 
Ontario, was discovered in 1995, and several thousand 
plants were noted at that time (Oldham, personal com- 
munication 2001). Surveys in 2001 documented approx- 
imately 1800 plants. The long sandspit island provides 
large areas of suitable habitat in Sable Island Prov- 
incial Nature Reserve. 

As a consequence of the ephemeral nature of its 
habitat and its ability to remain dormant in unfavour- 
able years, Lipocarpha micrantha may persist un- 
detected at locations searched thoroughly by botanists. 
An unsuccessful search during highwater, when much 
potential habitat is temporarily unavailable, is there- 
fore not justification for declaring a population extir- 
pated. This was the case at Poundnet Bay on Rainy 
Lake, Ontario, in 2001, where water levels were well 
above average. No Lipocarpha micrantha plants were 
found there during 2001 A population of 75 plants 
was discovered at this location in 2000. When water 
levels return to average or below this area will provide 
habitat for Lipocarpha micrantha. It is known from 
several beaches on the Minnesota side of Rainy Lake. 

Lipocarpha micrantha was first documented on 
Osoyoos Lake, British Columbia, by Ceska and Ceska 
(1980). Of the two populations noted, one has been 
eradicated by shoreline development. The British Col- 
umbia Conservation Data Centre has been monitor- 
ing the second population since 1991. In 2001 a total 
of between 30 000 and 50 000 plants were invento- 
ried. This is the highest recorded population at that site, 
and it is the largest population in Canada. The high 
number is attributed to excellent growing conditions, 
and also to the higher search intensity in 2001. If con- 
ditions remain stable at this site the population should 
continue to thrive. However, local development plans 
at the site, including a casino and marina, would likely 
destroy about 50 to 60% of the remaining habitat. A 
third, small (20 plants over 2 m*) population was dis- 
covered on the shoreline of Okanogan Lake in 2002. 


Special Significance of the Species 
Lipocarpha micrantha has very specific habitat 
requirements: open, sandy shorelines, protected from 


strong waves, with limited competition from other 
plants. It is extremely sensitive to alteration of both 
water levels and shoreline structure. As such it may 
be an important indicator of wetland quality. 


Protection 

Lipocarpha micrantha has a NatureServe (2002*) 
global rank of G5 or “secure”. In the United States, 
Lipocarpha micrantha is listed as Endangered in 
Connecticut, Maryland, New Jersey, New York, and 
Pennsylvania, and it is listed as Threatened in Maine 
and Ohio (United States Department of Agriculture, 
Natural Resources Conservation Service 2002*). In 
Canada, Lipocarpha micrantha is ranked as N1 or 
“critically imperilled”, and is listed as Threatened in 
Ontario (NHIC, 2003*) and Endangered/Threatened 
in British Columbia (Douglas et al. 2002). 

Both extant Ontario populations of Lipocarpha 
micrantha are in a Provincial Nature Reserve and a 
Provincial Conservation Reserve. As such they are pro- 
tected from shoreline development, but are subject to 
water level regulation. The extant populations in British 
Columbia occur on an Indian Reserve (Osoyoos Lake 
Indian Reserve #1) and in a Provincial Park (Sun Oka 
Beach Provincial Park). There is currently no specific 
provincial rare species legislation in place for the pro- 
tection of endangered/threatened vascular plants in 
British Columbia. At the federal level the Species at 
Risk Act protects COSEWIC-listed plants on federal 
lands (which includes Indian Reserves). It also empow- 
ers the federal government to protect habitat outside 
federal lands if the province fails to protect listed spe- 
cies or their habitat; these mechanisms are discretionary. 


Evaluation of Status 

The status of Lipocarpha micrantha in Canada is 
precarious. The largest, apparently stable, population 
at Osoyoos Lake in British Columbia is threatened by 
planned development. Should this development proceed, 
approximately half of the remaining Lipocarpha mic- 
rantha plants in Canada will be physically destroyed, 
and the remaining habitat may no longer support the 
species. The two remaining Ontario populations are 


2004 


SMITH, DOUGLAS, AND HARRIS: SMALL-FLOWERED LIPOCARPHA 


183 


FiGure 3. Distribution of Lipocarpha micrantha in Ontario and Quebec. Sable Island is shown as a circle, Ponies: Bay as 


a triangle. Squares denote extirpated populations. 


relatively small, both in number and area, and there- 
fore are vulnerable to environmental disturbance. If 
development at any of the remaining populations pro- 
ceeds without consideration for this species it will very 
likely be extirpated from Canada. In particular, altera- 
tion of natural water level cycles could result in the 
destruction of the habitat required by Lipocarpha mic- 
rantha. The authors’ recommendation that Lipocarpha 
micrantha be uplisted to Endangered was accepted by 
COSEWIC in November 2002. 


Acknowledgments 

The authors gratefully acknowledge the following 
people for assistance with this report. Kara Brodribb, 
Wasyl Bakowsky, Dan Dufour, Rob Foster, Ron Hall, 
Erica Oberndorfer, Mike Oldham, Kathy Paige, Amy 
Tesselin, and Christine Thuring assisted with field sur- 
veys. Wasyl Bakowsky, Brian and Rose Klinkenberg, 
Jim Meeker, Mike Oldham, Tony Reznicek and David 
Szymanski provided location and habitat data. Erica 
Oberndorfer, Jenifer Penny and Shyanne Smith assist- 
ed with the production of maps and compilation of 
status data. Graham Rose (Big Creek Hunt Club) gra- 
ciously granted access to their properties for survey 
work. This paper is based on a status report submitted 
to the Committee On the Status of Endangered Wildlife 


in Canada (COSEWIC), September 2001. The distri- 
bution map for Ontario and Quebec, Figure 3, was 
provided by COSEWIC. Funding was provided by 
COSEWIC, the George Cedric Metcalf Foundation, 
Royal Botanical Gardens, and the British Columbia 
Conservation Data Centre. 


Documents Cited 

Harris, A. G., M. J. Oldham, R. F. Foster, and W. D. 
Bakowsky. 2000. Preliminary life science inventory of 
Rainy Lake, Ontario. Rainy Lake Conservancy, Rainy 
Lake, Ontario. 

Holmgren, P. K, N. H. Holmgren, and L. C. Barnett. 2003. 
Index Herbariorum [web application]. Updated October 1, 
2003. New York Botanical Garden, Bronx, New York. 
Available at: http://www.nybg.org/bsci/ih/ih.html. 

Natural Heritage Information Centre. 2003. Natural Heri- 
tage Information Centre Database (http://www.mnr.gov. 
on.ca/MNR/nhic/data.cfm). Natural Heritage Information 
Centre, Ontario Ministry of Natural Resources. 

NatureServe 2002. NatureServe Explorer: An online ency- 
clopedia of life [web application]. 2002. Version 1.6. Arling- 
ton, Virginia, USA: NatureServe. Available at http://www. 
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Received 13 November 2003 
Accepted 5 April 2004 


Immobilization of Clover-trapped White-tailed Deer, Odocoileus 
virginianus, with Medetomidine and Ketamine, and Antagonism with 
Atipamezole 


JosHUA J. MILLSPAUGH!, BRIAN E. WASHBURN!, TAMARA M. MEYER?, JEFF BERINGER?, and LONNIE P. 
HANSEN? 


‘Department of Fisheries and Wildlife Sciences, University of Missouri, 302 Anheuser-Busch Natural Resources Building, 
Columbia, Missouri 65211 USA 

*Missouri Department of Conservation, Conservation Research Center, 1110 South College Avenue, Columbia, Missouri 
65201 USA 


Millspaugh, Joshua J., Brian E. Washburn, Tamara M. Meyer, Jeff Beringer, and Lonnie P. Hansen. 2004. Immobilization of 
Clover-trapped White-tailed Deer, Odocoileus virginianus, with medetomidine and ketamine, and antagonism with 
atipamezole. Canadian Field-Naturalist 118(2): 185-190. 


We evaluated the effectiveness of immobilizing Clover-trapped White-tailed Deer (Odocoileus virginanus) with medeto- 
midine hydrochloride (HCl) and ketamine HCI during winter and summer by monitoring immobilization intervals and vital 
signs. In winter, we captured deer in Clover traps in | 4-ha research enclosure for relocation to another on-site enclosure (n = 5). 
In summer, we captured free-ranging deer in Clover traps to attach radio-collars (n = 4). We administered an estimated 0.055 
mg/kg medetomidine HCI and 2.5 mg/kg ketamine HCI to adult (> 1.5 years of age) deer and 0.06 mg/kg medetomidine HC] 
and 2.5 mg/kg ketamine HCl to subadult (< 1.5 years of age) deer. We used an intramuscular injection of atipamezole HC] as the 
antagonist at a rate of 0.275 mg/kg for adults and 0.3 mg/kg for subadults > 30 minutes post-induction. Mean induction time in 
winter was 11.2 minutes (SE = 2.5, range = 5.4 — 24.2) and 6.5 minutes (SE = 0.8, range = 6.2 — 7.5) in summer. After atipame- 
zole HCl injection, the mean time to walking was 17.1 minutes (SE = 3.5, range = 7.5 — 41.5 minutes) in winter and 11.3 minutes 
(SE = 3.8, range = 4.7 — 13.5) in summer. Rectal temperature was relatively constant throughout immobilization; however 
rectal temperatures of 5 deer (7 = 3 in winter; n = 2 in summer) exceeded 40°C, a sign of hyperthermia. Respiration rate and pulse 
rate peaked at about 20 minutes post-medetomidine HCI and ketamine HCl injection, then generally declined thereafter. No 
mortalities were observed in our study. Medetomidine HCI and ketamine HCI doses for Clover-trapped White-tailed Deer pro- 
vided satisfactory induction times, sufficient level of anesthesia for short-distance relocation or radio-collar Eucte alee and 
were effectively reversed with an IM injection of atipamezole HCl. 


Key words: White-tailed Deer, Odocoileus virginianus, atipamezole, capture, chemical restraint, Clover trap, deer, ketamine, 
immobilization, medetomidine, Missouri. 


White-tailed Deer (Odocoileus virginianus) have 
been chemically immobilized with Telazol® (1:1 tile- 
tamine hydrochloride (HCl) and zolazepam HCl) and 
xylazine HCI (Schultz et al. 1992; Kilpatrick and Spohr 
1999), ketamine HCl and xylazine HCI (Mech et al. 
1985; Farley et al. 1986; Kreeger et al. 1986; Ballard et 
al. 1998; Kilpatrick and Spohr 1999), etorphine and 
xylazine HC] (Presnell et al. 1973; Presidente et al. 
1973; Nielsen 1982), xylazine HCI alone (Gibson et 
al. 1982), phencyclidine HCl (Dean et al. 1973), and 
succinylcholine chloride (Wesson et al. 1974; Jacobsen 
et al. 1976). Kreeger (1996) recommended 4.4 mg/kg 
Telazol® and 2.2 mg/kg xylazine HCl to immobilize 
White-tailed Deer; 0.125 mg/kg yohimbine HCl was 
the recommended antagonist. Recently, Kilpatrick and 
Spohr (1999) used a 4.8:3.9 mg/kg dose of Telazol®: 
xylazine HCI to dart free-ranging White-tailed Deer. 
Alternative drugs recommended by Kreeger (1996) 
include ketamine HCl and xylazine HCl, etorphine, 
xylazine alone (for calm deer only), and a combina- 
tion of medetomidine HCI and ketamine HCl. 

Medetomidine HCl depresses the central nervous 
system and acts similarly to xylazine HC] (Jalanka and 


Roeken 1990), but with greater affinity to alpha,- 
adrenoreceptors (Klein and Klide 1989; Jalanka and 
Roeken 1990; Kreeger 1996). Ketamine HCl, an anes- 
thetic, is often combined with a tranquilizer or seda- 
tive to improve induction and recovery (Haigh 1982; 
Kreeger 1996). Medetomidine HCl and ketamine HC] 
alone or in combination with other drugs have been suc- 
cessfully used to immobilize a diversity of large ungu- 
lates including Reindeer (Rangifer tarandus tarandus) 
(Ryeng et al. 2001, 2002), Mule Deer (Odocoileus 
hemionus) (Caulkett et al. 2000), Mule Deer/White- 
tailed Deer hybrids (Caulkett et al. 2000), Sika Deer 
(Cervus nippon) (Tsuruga et al. 1999), gemsbok (Oryx 
gazella) (Grobler et al. 2001), Roan Antelope (Hippo- 
tragus equinus) (Citino et al. 2001), Blue Duiker 
(Cephalophus monticola) (Bailey et al. 1995), Moose 
(Alces alces) (Arnemo 1995), Tigers (Panthera tigris) 
(Miller et al. 2003), Impala (Aepyceros melampus) (Bush 
et al. 2004), European Mink (Mustela lutreola) Four- 
nier-Chambrillon et al. 2003) and Red Deer (Cervus 
elaphus) (Arnemo et al. 1994). 

Notable among the advantages listed in these and 
other studies is the ability to reverse medetomidine HC] 


185 


186 


with an intramuscular (IM) injection of atipamezole 
HCl (Tsuruga et al. 1999; Haulena et al. 2000). Atipa- 
mezole HCI is an extremely efficient alpha,-adren- 
ergic antagonist compared with yohimbine HCl and 
tolazoline HCl (Kreeger 1996) and effectively reverses 
medetomidine HCl in many wildlife species (Tsuruga 
et al. 1999; Haulena et al. 2000; Grobler et al. 2001). 
If medetomidine HCl and ketamine HC] were as effi- 
cient and safe as other immobilizing agents (e.g., Tela- 
zol® and xylazine HCl) and if an IM injection of ati- 
pamezole HCI was a safe and effective antagonist, this 
combination could prove efficacious in field studies. 
Our objective was to determine the effectiveness and 
safety of immobilizing Clover-trapped White-tailed 
Deer with medetomidine HCI and ketamine HCI, and 
the feasibility of reversing this combination with an IM 
injection of atipamezole HCl, during winter and sum- 
mer by monitoring immobilization intervals and vital 
signs. To our knowledge, an evaluation using mede- 
tomidine HCI and ketamine HCl to immobilize free- 
ranging White-tailed Deer and antagonism by ati- 
pamezole HCI has not been previously published. 


Methods 

Our review of the medetomidine HCI and ketamine 
HCl combination took place in winter and summer 
2001. In summer and winter, we trapped White-tailed 
Deer in Clover traps (McCullough 1975). During win- 
ter we trapped deer at the Charles W. Green Conser- 
vation Area, located near Ashland, Missouri. Traps 
were baited with corn, set each evening, and checked 
at sunrise. All winter-trapped and immobilized deer 
(n = 5) were captured within a 4-ha research enclo- 
sure for relocation to another on-site 4-ha enclosure. 
These deer were not habituated to humans. 

In summer, we trapped White-tailed Deer at the 
Thomas S. Baskett Wildlife Research and Education 
Area, located near Ashland, Missouri. Traps, baited 
with salt, alfalfa, and corn, were set each evening and 
checked at sunrise. We captured and immobilized 
summer-trapped deer (n = 4) for purposes of radio- 
collar attachment. These deer were free-ranging ani- 
mals and not habituated to humans. 

In winter and summer, we immobilized adult deer 
(> 1.5 years of age) (n = 2 in winter; n = 3 in sum- 
mer) using an estimated 0.055 mg/kg medetomidine 
HCl and 2.5 mg/kg ketamine HCI; 0.06 mg/kg mede- 
tomidine HCI and 2.5 mg/kg ketamine HCI was ad- 
ministered to subadult deer (< 1.5 years of age) (n = 3 
in winter; n = 1 in summer). Adult deer, estimated to 
be 60 kg, were given 3.3 mg of | mg/ml Domitor® 
(medetomidine HCl; Orion Corporation, Orion— 
Farmos, Espoo, Finland) and 150 mg of 100 mg/ml 
Ketaset® (ketamine HCl; Fort Dodge Laboratories, 
Inc., Fort Dodge, Iowa, USA). Subadult deer, estimat- 
ed to be 30 kg, were given 1.8 mg of 1 mg/ml mede- 
tomidine HCl and 75 mg of 100 mg/ml ketamine HCl. 
We injected drugs IM into the biceps femoris with a 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


hand syringe. Sex and age (adult or subadult) were 
recorded and during winter each deer was marked with 
a plastic cattle ear tag in one ear for later identification. 
In summer, deer were fitted with radio-transmitters. 

Following sedation in winter and summer, we ap- 
plied an ophthalmic ointment and blindfolded the deer. 
In winter, we relocated deer to a different on-site 4-ha 
enclosure. Two or three field assistants placed the sed- 
ated deer into the rear of a vehicle (range from 100 to 
300 m away), which was driven to the release enclo- 
sure (< | km driving distance). Each deer was carried 
into the enclosure (< 30 m away), and placed in a ster- 
nal recumbent position. 

We reversed the medetomidine HCI and ketamine 
HCI combination with an IM injection of Antisedan® 
(atipamezole HCI; Orion Corporation, Orion—Farmos, 
Espoo, Finland) into the biceps femoris with a hand 
syringe at a rate of 0.275 mg/kg (16.5 mg of 5 mg/ml 
atipamezole HCl) for adults and 0.3 mg/kg (9 mg of 
5 mg/ml atipamezole HCl) for subadults. We visually 
monitored all deer until they departed the area. 

During winter and summer, we attempted to monitor 
immobilization intervals and vital signs at 5-minute 
intervals. For all deer, we recorded time of medetomi- 
dine HCI and ketamine HCl injection, induction (time 
from injection to time animal was handled), atipame- 
zole HCl administration, “head up” (time when the 
animal first lifted its head), “standing” (time when 
the animal first stood up), and “walking” (time when 
the animal successfully departed the area). We also 
recorded respiration rate (breaths/minute), rectal tem- 
perature (°C), and pulse rate (beats/minute) at 5-min- 
ute intervals beginning at the time of medetomidine 
HCI and ketamine HCl injection for respiration rate 
and beginning 10 minutes post-medetomidine HCl and 
ketamine HCl injection for temperature and heart rate. 


Results 

Nine deer (1 = 5 in winter, including 2 female sub- 
adults, 1 male subadult, | female adult, and 1 male 
adult; n = 4 in summer, including 3 adult females and 
1 female subadult) were immobilized using the drug 
combination described above and either relocated or 
equipped with a radio-collar. No mortality has been 
observed 10 months post-winter immobilization and 
5 months post-summer immobilization. 

Mean induction time was 11.2 minutes (SE = 2.5, 
range = 5.4 — 24.3) in winter and 6.5 minutes (SE = 0.8, 
range = 6.2 — 7.5) in summer. Time to atipamezole HCl 
injection averaged 54.4 minutes (SE = 3.7, range = 37.7 
— 79.2) in winter and 33.4 (SE = 1.3, range = 30.4 — 
33.8) in summer. After atipamezole HCl injection, the 
mean time to head up was 11.4 minutes (SE = 2.9, 
range = 4 — 27.5) in winter and 9.3 minutes (SE = 0.7, 
range = 8.4 — 9.9) in summer. Mean time to standing 
was 15.9 minutes (SE = 3.6, range = 4.4 — 41) in win- 
ter and 10.5 minutes (SE = 3.2, range = 4.7 — 12.3) in 
summer. The mean time to walking was 17.1 minutes 


2004 MILLSPAUGH, WASHBURN, MEYER, BERINGER, AND HANSEN: IMMOBILIZATION OF WHITE-TAILED DEER 187 


(SE = 3.5, range = 7.5 — 41.5 minutes) in winter and 
11.3 minutes (SE = 3.8, range = 4.7 — 13.5) in summer. 
With the exception of rectal temperatures, other 
vital signs were considered normal (Table 1). Rectal 
temperatures were stable from 10 — 30 minutes post- 
medetomidine HCl and ketamine HCl injection in 
summer and winter (Table |). Temperatures of 1 sub- 
adult female captured in winter were 37.4°C and 36.2°C 
at 70 and 110 minutes post-induction, respectively; 
thus, she was within 1.2°C of becoming hypothermic 
(defined as < 35°C; Kreeger 1996; DelGuidice et al. 
2001). No attempt was made to increase body temper- 
ature of this individual prior to antagonism with atipa- 
mezole HCl. After atipamezole HCI administration, 
it took that individual 41.5 minutes to depart, the max- 
imum time observed in our study. Rectal temperatures 
of 5 deer (n = 3 in winter, m = 2 in summer) exceeded 
40°C, a sign of hyperthermia. No attempt was made 
to decrease body temperature of these animals prior to 
antagonism. Respiration rates showed little variability 
in summer and winter and were generally in the upper 
20’s to low 30’s (breaths/minute) (Table 1) peaking 
at about 20 minutes post-medetomidine HCl and 
ketamine HCl injection. Pulse rates peaked about 20 
minutes post-medetomidine HCl and ketamine HCl 
injection at 95 beats/minute (SE = 3.6) in winter and 
103 beats/minute (SE = 1.4) in summer and declined 
thereafter to 74 beats/minute (SE = 5.6) in winter and 
88 beats/minute (SE = 3.1) in summer at 30 minutes 
post-medetomidine HCI and ketamine HCl injection 
(Table 1). No other adverse side effects were noted. 
During summer captures, response to IM injection 
of atipamezole HCl was predictable, as previously 
described by Jalanka and Roeken (1990). Within 3 — 
6 minutes of atipamezole HCl injection, “ear-twitch- 
ing” occurred, followed by leg extensions after an addi- 


tional 3 — 6 minutes, and “head up” 2 minutes there- 
after. Standing followed within another 2 minutes and 
the animal departed almost immediately with good 
muscle coordination. 


Discussion 

Medetomidine HCl] and ketamine HCl doses for 
Clover-trapped deer provided satisfactory induction 
times, produced a sufficient level of anesthesia for 
short-distance relocation or radio-collar attachment, 
and were effectively reversed with an IM injection of 
atipamezole HCI. Small dosage volume, ease of prepa- 
ration and predictable responses to sedation and to 
the antagonist make this combination a useful alterna- 
tive to drug combinations that may require prolonged 
recovery. 

Medetomidine HCl and ketamine HCl provided 
induction times similar to those reported in other 
studies and with other drugs. For 13 captive White- 
tailed Deer in Minnesota, it took 2 to 35 minutes (medi- 
an = 8, SE = 1.2) from the time of xylazine HCI and 
ketamine HCl administration before deer lost the ability 
to stand (Mech et al. 1985). Our mean induction time 
in winter (x = 11.2 minutes, SE = 2.5, range = 5.4 — 
24.3, n = 5) was about double the median time of 6.2 
minutes (range = 0.5 — 17.3) reported by Jalanka and 
Roeken (1990) for 28 captive White-tailed Deer housed 
at the Helsinki Zoo, but similar to our summer mean 
time of 6.5 minutes (SE = 0.8, range = 6.2 — 7.5). In 
the Helsinki Zoo study, deer were given an average 
of 61 ug/kg (SD = 14, median = 58, range = 37 — 98) 
medetomidine HCI and a mean ketamine HCI dose of 
1.6 mg/kg (SE = 0.3, median = 1.5, range = 1 — 2.3). 

The time to walking after the IM injection of ati- 
pamezole HCI was similar to that for deer in other 
studies reversed with yohimbine HCl, but was less 


TABLE |. Mean + SE (N) vital signs of White-tailed Deer immobilized with medetomidine hydrochloride (HCl) and ketamine 
HCl, and antagonized with atipamezole HCl during winter and summer 2001 in mid-Missouri for purposes of relocation 
(winter) and radio-collar attachment (summer). Adults were immobilized with 0.055 mg/kg medetomidine HCI and 2.5 
mg/kg ketamine HCl and subadults were immobilized with 0.06 mg/kg medetomidine HCl and 2.5 mg/kg ketamine HCl. 
Adults were antagonized with 0.275 mg/kg of atipamezole HCI and subadults were antagonized with 0.03 mg/kg of atipamezole 
HCI. We have not reported data where information was collected on < 2 deer. Time 0 is the time of medetomidine HCI and 


ketamine HCl injection. 


Winter Summer 
Time Respiration Rate Pulse Rate Temperature Respiration Rate Pulse Rate Temperature 
(minutes) (breaths/minute) (beats/minute) (c) (breaths/minute) (beats/minute) (°C) 
0 PH pd =e PEMA G)) — - 23 + 1.4 (4) - ~ 
5 DifeD) 7D) -3)()) - - 23 + 2.2 (4) - - 
10 28.8 + 2.9 (5) 87 +3.6(4) 41.1405 (2) 31+2.4 (4) 98+2.0(4) 39.9+ 1.2 (4) 
15 29.6 + 3.2 (5) 88 +43(4) 40.9+0.8 (4) 31+2.2(4) 101+1.9(4) 39.8414 (4) 
20 SI =n 244) 94.7+3.6(3) 40.62 1.2 (3) 30+2.6(4) 103+1.9(4) 39.8+1.4(4) 
25 Posy \oe2 Syil (G) 86 +3.5(4) 404+1.2 (4) 30+3.1(4) 101424(4) 39.6+1.3 (4) 
30 29.3 + 3.6 (4) Ase) 16\(2) wae 3 9-3 -t ile 9 (2) 32 + 2.9 (4) 88+3.1(4) 39.3+1.2 (4) 
35 24 +2.4(2) Diet) ED (2) - 24 + 0.0 (2) ~ = 
40 - — ~ 24 + 0.0 (2) - ~ 
45 Syd" Tes 3}5) (3) - - — — = 
50 PR: a2 3)3} (8) - 40.1 + 1.2 (2) - - - 


188 


variable. An intravenous injection of yohimbine HCl 
following xylazine HCl and ketamine HCl immo- 
bilization resulted in an adult male walking in 1.5 min- 
utes (SE = 0.5, n = 2 immobilizations) to 26.5 minutes 
(SE = 11.5, n = 4 immobilizations) for an adult fe- 
male (Mech et al. 1985). For 22 White-tailed Deer 
(18 free-ranging captured using drop nets and 4 cap- 
tive) immobilized with xylazine HCl and ketamine 
HCl, an IM injection of yohimbine HCI produced a 
mean recovery time of 11.6 minutes (SE = 2.3) 
(Wallingford et al. 1996). Hsu and Shulaw (1984) used 
an IV injection of yohimbine HCI and reported a mean 
recovery time of 4.4 minutes (SD = 5.4 minutes) for 
xylazine HCl-immobilized deer. 

With the exception of rectal temperature, the vital 
signs observed in this study were within the normal 
range of reported values for White-tailed Deer. Mautz 
and Fair (1980) reported pulse rates of a 46-kg adult 
White-tailed Deer female in July that was lying, stand- 
ing/walking, and running at 65, 74, and 106 beats per 
minute, respectively. Pulse rates observed in this study 
were similar to predicted walking pulse rates of White- 
tailed Deer (Moen 1978: 722), yet higher than those 
rates reported by Jalanka and Roeken (1990: 267). 
Rectal temperatures of five deer were above 40°C, a 
sign of hyperthermia (Kreeger 1996). Average rectal 
temperatures in our study in winter (39.3°C at 30 
minutes, SE = 1.9, n = 2) and summer (39.3°C at 30 
minutes, SE = 1.2, n = 4) are similar to the maximum 
rectal temperatures reported by DelGuidice et al. 
(2001:1151) for White-tailed Deer captured by Clover 
trap and immobilized with xylazine HCI and ketamine 
HCI during the winter in Minnesota. Also in Minne- 
sota, Rogers et al. (1987) reported rectal temperatures 
of two female free-ranging White-tailed Deer fawns in 
all seasons during various activities ranged between 
38.2° and 40.1°C. Rectal temperatures of 3 adult male 
White-tailed Deer in Mississippi, averaged 39.3°C 
during late August and September and 38.6°C in 
early December (Demarais et al. 1986). As suggested 
by DelGiudice et al. (2001), corrective actions (i.e., 
packing snow around the animal) should be taken 
when immobilized animals become hyperthermic. 
Consequently, rectal temperatures should be monitored 
and protocols should be established to determine when 
corrective actions should begin (DelGiudice et al. 
2001). 

Both medetomidine HCl and ketamine HCl have 
wide safety margins, produce calm inductions in several 
artiodactyls, are safe for humans (Jalanka and Roeken 
1990), and have not caused any apparent detrimental 
effects in pregnant females (Jalanka 1993). Few adverse 
side effects were noted by Jalanka and Roeken (1990) 
after 1240 immobilizations with medetomidine HCl, 
ketamine HCl, and antagonism with atipamezole HCl. 
Worth noting, some ruminants became resedated be- 
tween 30 and 240 minutes post-IV reversal with ati- 
pamezole HCI (Jalanka and Roeken 1990) and unre- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


markable ruminal tympany was common in ruminants 
prior to atipamezole HCI administration (Jalanka and 
Roeken 1990; Jalanka 1993). 

A disadvantage of the medetomidine HCl, ketamine 
HCl, and atipamezole HC] combination is cost. In U.S. 
currency, cost to immobilize each adult deer was 
$34.72 ($32.76 for medetomidine HCI and $1.96 for 
ketamine HCl) and $18.85 for each subadult deer 
($17.87 for medetomidine HC] and $0.98 for ketamine 
HCl). Atipamezole HCI cost $33.22 per adult deer and 
$18.14 for each subadult deer. In contrast, Kilpatrick 
and Spohr (1999) reported a cost of $8.44/deer for 
Telazol® and xylazine HCl immoblized deer and $6.34/ 
deer for ketamine HCI and xylazine HCI without lyo- 
philizing costs. With lyophilizing, costs were $10.05/ 
deer for Telazol® and xylazine HCl and $16.49 for 
ketamine HCl and xylazine HCI (Kilpatrick and Spohr 
1999). Consequently, use of medetomidine HCl, keta- 
mine HCl, and atipamezole HCI may be cost-prohib- 
itive in studies requiring immobilization of many ani- 
mals. However, given the desirable properties discussed 
above, including rapid reversal that requires less man- 
power time, these desirable qualities may outweigh 
drug costs. Also, the ability to administer an antagonist 
IM may be advantageous in field studies (Wallingford 
et al. 1996). We recommend researchers investigate 
the utility of reversing the less costly and popular 
Telazol®/xylazine HCl and ketamine HCl/xylazine 
HCl combinations with an IM injection of atipame- 
zole HCl. 

The combination of medetomidine HCl, ketamine 
HCl, and atipamezole HCl reported herein provided 
an effective level of anesthesia for Clover-trapped 
White-tailed Deer. We recommend the use of 0.055 
mg/kg medetomidine and 2.5 mg/kg ketamine HCl to 
immobilize adult Clover-trapped White-tailed Deer 
and 0.06 mg/kg medetomidine HCl and 2.5 mg/kg 
ketamine HC] for subadults. Furthermore, we recom- 
mend 0.275 mg/kg and 0.3 mg/kg of atipamezole, 
injected IM, to reverse this combination in adults and 
subadults, respectively. 


Acknowledgments 

We thank R. Houf, T. Mong, and numerous volun- 
teers for their assistance in the field. Missouri Depart- 
ment of Conservation, Conservation Research Center, 
provided Clover traps. A University of Missouri Res- 
earch Board grant, the University of Missouri, and the 
Missouri Department of Conservation supported this 
project. 


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Arnemo, J. M., T. Negard, and N. E. Soli. 1994. Chemical 
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190 


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THE CANADIAN FIELD-NATURALIST 


Vol. 118 


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Received 2 March 2002 
Accepted 3 October 2004 


reer 


Predation on Two Mule Deer, Odocoileus hemionus, by a Canada Lynx, 
Lynx canadensis, in the Southern Canadian Rocky Mountains 


Dore Poszic!, CLAYTON D. Apps?, and ALAN Disb? 


' Alhardstr. 15, 28757 Bremen, Germany 


? Aspen Wildlife Research Inc., 2708 Cochrane Road N.W., Calgary, Alberta T2M 4H9 Canada (corresponding author) 
3 Parks Canada, P. O. Box 220, Radium Hot Springs, British Columbia VOA 1MO0 Canada 


Poszig, Dérte, Clayton D. Apps, and Alan Dibb. 2004. Predation on Two Mule Deer, Odocoileus hemionus, by a Canada 
Lynx, Lynx canadensis, in the southern Canadian Rocky Mountains. Canadian Field-Naturalist 118(2): 191-194. 


A male Canada Lynx (Lynx canadensis) killed two Mule Deer (Odocoileus hemionus) in the southern Canadian Rocky 
Mountains in January 1999 and made use of the kills for 28 days. Canada Lynx predation on ungulates has been reported but 
is rare, and accounts have been brief. We detail the lynx behaviour associated with the kills and their consumption. An infra- 
red monitor and attached camera were used to register daily activity at the kill site. We speculate on the factors that may have 


influenced this opportunistic predation event. 


Key Words: Canada Lynx, Lynx canadensis, infrared monitor, Mule Deer, Odocoileus hemionus, predation, British Columbia. 


The Canada Lynx (Lynx canadensis), hereafter 
referred to as lynx, preys mainly on Snowshoe Hares 
(Lepus americanus) and other small mammals, but 
rare accounts of ungulate predation have been noted 
(e.g., Saunders 1963; van Zyll de Jong 1966; Parker 
et al. 1983; Stephenson et al. 1991). However, these 
accounts do not describe lynx behaviour associated 
with the kills or their consumption. We give a detailed 
description of two Mule Deer (Odocoileus hemionus) 
kills made by a lynx and its subsequent use of them 
for 28 days. The observations were made in the course 
of an in-depth field study of lynx ecology in the south- 
ern Canadian Rocky Mountains of southeastern British 
Columbia and southwestern Alberta. 

While snow-tracking on 18 January 1999, we found 
two Mule Deer that had been killed by a radio-col- 
lared adult male Canada Lynx in Kootenay National 
Park, British Columbia (51° N, 116° W). One was a 
doe aged by cementum annuli to be 3.5 years, and we 
estimated the other deer to be a fawn (teeth not com- 
pletely erupted). The kills were separated by a distance 
of 10 m and were located at 1580 m elevation on a 
15° slope of northwest aspect in a closed-canopy forest 
of Lodgepole Pine (Pinus contorta), Engelmann Spruce 
(Picea engelmannii), and Subalpine Fir (Abies lasio- 
carpa). The understory was sparse, although the imme- 
diate area around the kill site had some woody debris 
to a depth of about | m and some conifer thickets. The 
snow depth at the site was 90 cm and lynx penetration 
was 9-14 cm. 

The fawn apparently was killed on 14-15 January, 
judging from daily radio-telemetry fixes of the lynx 
and accumulated snow cover. The shoulders, rib cage, 
part of the neck, upper front legs and organs had been 
consumed. The front and rear had been severed and lay 
twisted and partially cached in a hollow in the snow. 
The doe was killed on 18 January between about 1300 


and 1500 h. On initial inspection, no snow had accu- 
mulated on the carcass, and the blood had not yet co- 
agulated or frozen. The deer was found lying on its side 
with legs extended, and no part had yet been consumed. 

Teeth of both deer were in good condition, and no 
deformities of their legs or hooves were apparent. The 
doe appeared to be in good physical condition. Both 
deer had major wounds on the dorsal side of their necks, 
and the spacing of obvious tooth-punctures matched 
that of a lynx (Figure 1). The cause of death of the 
fawn could not be determined because it had been con- 
sumed to a great extent and was not intact. On the doe, 
several puncture wounds were located in two small 
areas on the dorsal to slightly lateral area of the neck, 
10-15 cm behind the ears. Claw marks on the deer’s 
back and shoulders suggested that the lynx had leapt 
onto the deer and had been “riding” while biting it. 

Evidence of predation of the fawn was obscured by 
recent snow, but old blood stains 2 m from the carcass 
suggested that it had been killed and not scavenged. 
When removing the mandible of the fawn, we found 
pieces of undigested Subalpine Fir in its mouth, sug- 
gesting that the animal had been browsing shortly 
before it died. 

Tracks in the snow provided a clear record of how 
the lynx encountered and stalked the doe. The lynx 
approached this deer from a slightly higher elevation 
than where both kills were found, partially circling the 
site for about 100 m. It briefly stopped at the edge of 
a small opening about 50 m from the kill site, and then 
again 25 m uphill on a knoll. Moving downhill in a 
normal gait, the lynx used several fallen logs for cover, 
but was clearly stalking for the last 10 m. It then made 
three bounds before its attack, as the doe stood next 
to a thicket of mixed-age trees. The deer appeared to 
succumb <4 m from the attack site. The lynx then 
dragged the carcass about 7 m to its final location. 


LOM 


192 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


FiGure 1. Neck wound (arrows indicate puncture locations) resulting in the death of an adult female Mule Deer by a 
Canada Lynx in the southern Canadian Rocky Mountains, January 1999. 


Six site visits conducted over the three weeks fol- 
lowing the initial find indicated that the lynx first fed 
on the doe’s neck, continued to feed on the shoulders 
and rib cage, and finally ate the rear and internal organs. 
Deer hair had been removed and was positioned around 
and on top of the kill. There were several well-used 
lynx beds that appeared to be strategically located 
upslope, <120 m from the kill site, with clear down- 
hill views. The hindquarters of the fawn were moved 
to 4 different locations between our site visits, and 
each time were cached under snow and branches. It is 
possible that this was prompted by our inspections. 

On 29 January we installed an infrared monitor 
(Trailmaster™ 1500, Goodson and Associates Inc., 
Kansas, with attached Olympus camera) at the doe’s 
kill site to document the 24-hour activity of the lynx 
at the site. The monitor was programmed to register 
all beam interferences, while the remote camera was 
set to take a photo upon beam interference, with <1 
photo/15 min and between 0900 h and 1630 h to avoid 
flash disturbance. Because the lynx no longer appeared 
to be feeding on the fawn, we expected that all subse- 
quent feeding events would be registered. 

Twenty photos of the lynx were obtained and 
showed that only this animal fed on the carcass (e.g., 
Figure 2). In addition, no other predator was detected 
at the site during inspections that occurred about every 
four days. lynx activity at the kill peaked around mid- 


night and in the late morning (Figure 3). 

The lynx abandoned the kill site after 28 days, at 
which time the head and lower side of the doe’s car- 
cass were frozen into the ground. Scratch marks sug- 
gested that the lynx had unsuccessfully tried to remove 
some snow to access parts of the deer. Upon incidental 
recapture 16 days later, the lynx weighed 15.9 kg, 
18.6% more than at his previous and original capture 
in November 1996, while skeletal measurements had 
not changed (C. D. Apps, unpublished data). 

Our account is consistent with the observations by 
Stephenson et al. (1991) of lynx ambushing ungulate 
prey at close range. They reported a lynx remaining 
on a Caribou (Rangifer tarandus) kill for 42 days. In 
our study, the Lynx apparently abandoned the site be- 
cause it could not make use of the carcass remains, and 
it might have stayed longer had conditions permitted. 

Previous reports of lynx predation on ungulates 
have been associated with the low phase of the approx- 
imate 10-year population cycle of Snowshoe Hares, 
the lynx’s primary prey. The reliance of lynx on alter- 
nate prey is greatest at this time (Mowat et al. 2000). 
Although the deer kills we report occurred during an 
assumed Snowshoe Hare population peak, hare densi- 
ties in the study area were comparable to boreal regions 
during the cyclic low (Apps 2000; C.D. Apps, unpub- 
lished data). 


2004 Poszic, APPS, AND DIBB: PREDATION ON MULE DEER By A LYNX 193 


= 


itl 


FIGURE 2. Remote camera photo of a Canada Lynx at the carcass of a Mule Deer it killed in the southern Canadian Rocky 
Mountains, January 1999. 


80 10 
a Frequency of Monitor interference 
~9 
70 . 
=—@—Frequency of Days with > 0 
Monitor Interferences 8 
60 
Uf 
50 
6 
3 g 
S 40+ 5 s 
S a 
we 4 
30 
3 
20 
2 


10 


00:00- 02:00- 04:00- 06:00- 08:00- 10:00- 12:00- 14:00- 16:00- 18:00- 20:00- 22:00- 
01:59 03:59 05:59 07:59 09:59 11:59 13:59 15:59 17:59 19:59 21:59 23:59 


Time of day (h) 


Ficure 3. Lynx activity while feeding on a Mule Deer kill over 11 days in the southern Canadian Rocky Mountains, 
January 1999. Total frequency (1 min. intervals; n = 344) and number of days that the Canada Lynx triggered an infrared 
monitor (monitor interference) per 2-h period are shown. 


194 


We further note that the relatively high elevation 
and deep snow of the kill site made it an atypical loca- 
tion for Mule Deer during mid-winter. We speculate 
that the deer may have been avoiding other predators, 
most notably Gray Wolves (Canis lupus) that travel 
primarily through the valley bottom. Thus, we expect 
that this predation was opportunistic, perhaps influ- 
enced by relatively low densities of Snowshoe Hares, 
the age class and sex of the lynx (adult male), and the 
vulnerability of the deer. 


Acknowledgments 

The Southern Canadian Rockies Lynx Project was 
jointly funded by the Habitat Conservation Trust Fund 
through the British Columbia Ministry of Water, Land 
and Air Protection (formerly Environment, Lands 
and Parks), Parks Canada, and the Columbia Basin 
Fish and Wildlife Compensation Program. Additional 
financial contributions were provided by the Friends 
of Kootenay National Park, Husky Oil, World Wild- 
life Fund Canada, Fairmont Hotels and Resorts (for- 
merly Canadian Pacific Hotels & Resorts), and the 
Bow Valley Naturalists. Contract supervision, input, 
and direction were provided by A. Dibb of Parks Can- 
ada, J. Krebs of the Columbia Basin Fish and Wildlife 
Compensation Program, and A. Fontana, R. Neil, G. 
Tipper, R. Forbes, and I. Teske of BC Environment. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Literature Cited 

Apps, C. D. 2000. Space use, diet, demographics, and topo- 
graphic associations of Lynx in the southern Canadian 
Rocky Mountains. Pages 351-371 in Ecology and conser- 
vation of Lynx in the United States. Edited by L. F. Rug- 
giero, K. B. Aubry, S. W. Buskirk, G. M. Koehler, C. J. 
Krebs, K. S. McKelvey, and J. R. Squires. University Press 
of Colorado, Boulder, Colorado. 

Mowat, G., K. G. Poole, and M. O’Donoghue. 2000. 
Ecology of Lynx in northern Canada and Alaska. Pages 
365-306 in Ecology and conservation of Lynx in the 
United States. Edited by L. F. Ruggiero, K. B. Aubry, S. 
W. Buskirk, G. M. Koehler, C. J. Krebs, K. S. McKelvey, 
and J. R. Squires. University Press of Colorado, Boulder, 
Colorado. 

Parker, G. R., J. W. Maxwell, L. D. Morton, and G. E. J. 
Smith. 1983. The ecology of the Lynx (Lynx canadensis) 
on Cape Breton Island. Canadian Journal of Zoology 61: 
770-786. 

Saunders, J. K. Jr. 1963. Food habits of the Lynx in New- 
foundland. Journal of Wildlife Management 27: 384-390. 

Stephenson, R. O., D. V. Grangaard, and J. Burch. 1991. 
Lynx, Felis lynx, predation on Red Foxes, Vulpes vulpes, 
Caribou, Rangifer tarandus, and Dall Sheep, Ovis dalli, 
in Alaska. Canadian Field-Naturalist 105: 255-262. 

van Zyll de Jong, C. G. 1966. Food habits of the Lynx in 
Alberta and the Mackenzie District, N.W.T. Canadian 
Field-Naturalist 80: 18-23. 


Received 8 September 2002 
Accepted 25 October 2004 


Effect of Fire Intensity and Depth of Burn on Lowbush Blueberry, 
Vaccinium angustifolium, and Velvet Leaf Blueberry, Vaccinium 
myrtilloides, Production in Eastern Ontario 


Luc C. DUCHESNE! and SUZANNE WETZEL2? 


'Rorest BioProducts Inc., 945 Queen Street East, Sault Ste Marie, Ontario P6A 2E5 Canada 
>Natural Resources Canada, Canadian Forestry Service, 1219 Queen Street East, Sault Ste Marie, Ontario P6A 3A5 Canada 
3Corresponding author at swetzel@NRCan.gc.ca 


Duchesne, Luc C., and Suzanne Wetzel. 2004. Effect of fire intensity and depth of burn on Lowbush Blueberry, Vaccinium angusti- 
folium, and Velvet Leaf Blueberry, Vaccinium myrtilloides, production in eastern Ontario. Canadian Field-Naturalist 
118(2): 195-200. 


The effects of prescribed fire intensity and depth of burn were investigated on Lowbush Blueberry (Vaccinium angustifolium) 
and Velvet Leaf Blueberry (Vaccinium myrtilloides) stem density, blueberry production and the number of blueberries/stem in a 
clear-cut Jack Pine, Pinus banksiana, ecosystem of eastern Ontario. Blueberry production and stem density were significantly 
(P < 0.001) increased by low intensity prescribed fires of 597 and 1268 kW/m. In contrast, prescribed fires of medium and high 
intensities did not affect blueberry production and stem density. The number of blueberries/stem was not affected (P = 0.056) 
by prescribed burning, two years after treatment. Pearson’s multiple correlation analysis showed that blueberry production 
(R: -0.683, P < 0.01), stem density (R: 0.733, P < 0.01) and the number of blueberries/stem (R: 0.803, P < 0.01) correlated with 
depth of burn. As well, blueberry production (R: 0.507, P < 0.05) and stem density (R: -0.504, P < 0.05) correlated with fire 
intensity. Depth of burn was a better predictor of berry production and stem density than fire intensity. These results suggest 
that only low intensity fires with little penetrating effect in the ground should be used to manage blueberry crops. 


Key Words: Lowbush blueberry, Vaccinium augustifolium, Vaccinium myrtilloides, wildfire, nontimber forest products (NTFP). 


Nontimber forest products (NTFP) are all botanical 
commodities harvested from the forest excluding in- 
dustrial timber use (Duchesne et al. 2000). NTFP are 
critical to the economy of forest and rural communi- 
ties by providing food as well as supplemental income, 
which often improves the standard of living of rural 
and First Nation communities (Brubaker 1997; Du- 
chesne et al. 2001; Mohammed 1999). Despite the 
socio-economic importance of this industry there has 
been little scientific research conducted for its support, 
presumably because of its novelty. However, the long- 
term success of the NTFP industry depends on acquir- 
ing precise knowledge regarding the availability, and 
sustainability of wild harvests as well as strategies to 
domesticate and manage many of the most sought after 
NTFP species. Species such as Lowbush Blueberry 
(Vaccinium angustifolium) and Velvet Leaf Blueberry 
(Vaccinium myrtilloides) need to be further domesti- 
cated and/or managed in natural ecosystems to meet 
the growing demand from the food and nutraceutical 
industries (Duchesne et al. 2001). This paper describes 
research conducted to improve blueberry management. 

Canada is the world’s largest producer of wild blue- 
berries (Lowbush and Velvet Leaf blueberries) with 
an annual output of $42 425 million in farm gate value 
resulting in production of 43 511 tonnes of frozen 
blueberries (Agriculture and Agri-Food Canada 2002). 
The demand for Lowbush and Velvet Leaf blueberries 
is such that management of natural or established sites 
is conducted to promote optimal yields ranging from 
3360 to 8967 kg/ha (Mohammed 1999). Given that 


blueberries are worth approximately $1.00/kg to farm- 
ers, the annual output of managed fields ranges from 
$3360 to $8967/ha, which is far superior to the aver- 
age timber growth of Canada’s forests of 1.59 m°/ha/ 
year (Lowe et al. 1996) with an approximate value of 
$100/ha/year. 

Pruning of blueberry fields, either by mowing or 
burning, is used routinely to maximize blueberry pro- 
duction (Badcock 1958; Eaton 1958; Eaton and White 
1960) as it stimulates the development of new shoots 
from shallow rhizomes (Van Hoefs and Shay 1981). In 
turn, new shoots are preferred over old shoots as the 
ratio of flower buds to leaf buds is greater on new 
shoots than on older shoots (Hall et al. 1972). Fire 
pruning of blueberry fields is generally accomplished 
by using straw as a fire carrier or tractor mounted oil or 
propane gas burners that emit a constant flame (Blatt 
et al. 1989). Burning also has the added advantage of 
reducing problems with insects and disease (Blatt et 
al. 1989; Smith and Hilton 1971). Although fire is 
frequently used for the management of Lowbush and 
Velvet Leaf blueberries (Blatt et al. 1989), which form 
one commercial crop, there is a lack of information 
about the relationship between fire behaviour and its 
impact on berry production. Indeed, current prescrip- 
tions aim at using fires that kill all stems (Blatt et al. 
1989). However, fire behaviour varies a great deal even 
during prescribed burns, which in turn affects the eco- 
logical outcome of post-fire ecosystems (McRae et al. 
2001). 


195 


196 


Frontal fire intensity (or Byram’s fireline intensity) 
is a measurement of the quantity of energy liberated as 
a fireline moves through an ecosystem, and is the most 
encompassing physical measurement of fire behavior 
related to ecological impacts (reviewed by Alexander 
1982 and McRae et al. 2001). Fire intensity is some- 
times used together with depth of burn (Miller 1977), 
which is a measurement of heat penetration into the 
soil. However, both frontal fire intensity and depth of 
burn are relatively new concepts in fire ecology and 
have never been applied to blueberry management. 
Hence, the objective of this study was to investigate 
the effect of fires of different intensities and depth of 
burn on Lowbush and Velvet Leaf blueberry pro- 
duction and vegetative growth. 


Materials and Methods 
Study site 

The study area is located at Frontier Lake (46°00’N, 
77°33’ W) in a Jack Pine, Pinus banksiana, stand in 
eastern Ontario within the middle Ottawa section (L.4c) 
of the Great Lakes — St. Lawrence Forest region 
(Rowe 1972). As reported by Herr et al. (1994), the 
site is near the Petawawa Research Forest, and is rela- 
tively flat, with a difference in elevation of approx- 
imately 4 m over 1.0 km. The surface deposit is a 
fine-grained and deep sand (10-30 m deep) (Gadd 
1962) and the soil is a humo-ferric podzol. The study 
site was selected because of its uniformity in tree com- 
position and topography. It was harvested in 1942 
and 1943 leaving residual standing timber with a stump 
diameter of 17.5 cm or less. Dendrochronological 
analysis of dominant trees and snags with multiple 
fire scars suggests that the study site sustained sev- 
eral fires, with the most recent in 1943, presumably 
from broadcast slash burning following harvesting 
(E. Stechishen, personal communication). 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Presently, the stand comprises a mix of Jack Pine 
(Pinus banksiana), Red Pine (Pinus resinosa) and 
White Pine (Pinus strobus) with the Red and White 
pine forming an emergent layer (Herr et al. 1994). 
Although the Jack and Red pine are of similar ages, 
the Jack Pine has higher relative density and is the 
dominant tree species on the site. Other plant species 
found at the site include serviceberry (Amelanchier 
sp.), Sweet Fern (Comptonia peregrina), Wintergreen 
(Gaultheria procumbens), Sheep Laurel (Kalmia an- 
gustifolia), Ground Pine (Lycopodium complanatum), 
Cow Parsnip (Maianthemum canadense), Hairy Solo- 
mon’s Seal (Polygonatum pubescens), Bracken Fern 
(Pteridium aquilinum ) and Sand Cherry (Prunus 
pumila). 


Treatments 

In the summer of 1990, an area of 150 m x 1000 m 
was clear-cut with the slash left in place. The site was 
divided into 40 plots of 35 m x 70 m with 8 m fire- 
guards established around each plot. In 1991, ten plots 
were burned under different indices of the Canadian 
Forest Fire Weather Index (FWI) system (Table 1) 
resulting in frontal fire intensities that varied from 
597 to 21 305 kW/m and depth of burn that varied 
from 0.37 to 2.78 cm (McAlpine 1995). This range 
of fire intensity and depth of burn far exceeds the 
range of fire behaviour currently used in blueberry 
management (unpublished observations). 

Pre- and post-burn fuel loads were measured for 
both slash and duff fuels (McRae et al. 1979). Fire rate 
of spread was measured with a pin grid network on 
each plot; fire arrival times at each pin were recorded 
to provide distance and time information. Fuel con- 
sumption was determined as the difference between 
these two values. Fire intensity was calculated using 
Byram’s (Byram 1959: cited in McAlpine 1995) 


TABLE |. Fire weather', fire behaviour parameters! and blueberry production from thirteen 35 m x 75 m plots in a clear cut 
Jack Pine after prescribed burning of different fire intensities at Frontier Lake Experimental site in Eastern Ontario. 


Plot Date Depth! of Intensity! Blueberries” Stem density* Blueberries/stem* 
Number of burn! FWI! — Burn (cm) (kW/m) (g/m?) (stems/m?) (N/plant) 

2 08/08/91 3.1 0.37 597 492.86 a (167.4) 73.2 a (8.5) 7.53 a (3.4) 

7 08/08/91 3.6 0.38 1 268 242.18 a (79.44) 44.2 b (9.8) 5.52 a (1.6) 
44 06/24/91 15.8 1.79 2 305 10.81 b (7.6) 5.9 c (0.7) 2.26 a (1.8) 
40 07/10/91 4.3 2.78 4 844 114 b, (©@:9) 9.4 c¢ (2.9) 0.90 a (0.69) 
32 07/12/91 9.6 1.85 7 600 8.78 b (8.7) 6.7 c (5.6) 0.53 a (0.46) 
26 06/14/91 11.0 3.50 10 941 4.96b (4.8) 9.9 ¢ (7.1) 3.40 a (2.9) 
46 06/24/91 17.4 1.95 12 065 ZO Mb Gels) 4.0 c (1.9) 3.10 a (2.7) 
36 07/12/91 3.9 Dail 13 202 27.20b (8.9) 12.1 c (4.6) 2.59 a (1.13) 
43 07/10/91 8.8 2.64 20 334 0.53b (0.24) 5.7 c (4.1) 0.50 a (0.42) 
24 06/14/91 21.0 1.79 21 305 5.33b (0.48) 13.8 c¢ (2.4) 0.04 a (0.06) 

Controls? — = — 9.01b (2.6) 5.9 c (1.48) 0.78 a (0.25) 


' from McAlpine (1995). 


* within column means followed by a different letter are significantly different at P < 0.05 determined by ANOVA. Observed 
significance level was adjusted with the Bonferroni procedure. 
> controls consisted of three unburned plots adjacent to the burned over plots. 


2004 


intensity equation I=HwR, where I is the intensity of 
the fire (kW/m), H is the fuel low heat of combustion 
(assumed to be 18 000 kJ/kg), w is the weight of fuel 
consumed in the active front (kg/m?) (all fuel con- 
sumed was assumed to have been burned by the active 
fire front), and R is the rate of spread (m/sec). 

To determine depth of burn, ten 50 cm steel pins 
mounted with horizontal markers were used in each 
plot to mark the top of the soil litter layer prior to each 
burn. Immediately after the burn, the distance between 
the horizontal marker and the top of the litter layer 
was measured with a ruler and called depth of burn. 

Two years following prescribed burning, there were 
45 plant species in our research plots. The most impor- 
tant species in terms the biomass and frequency (in 
decreasing order of importance) were P. aquilinum, 
V. angustifolium, V. myrtilloides, Amelanchier sp., P. 
pumila, Comptonia peregrina, Kalmia angustifolia, 
Yellow Panic-grass (Panicum xanthophysum), and 
Houton’s Sedge (Carex houghtonii) (Tellier et al. 1995, 
1996). 


Blueberry collection 

To compare the effect of fire intensity and depth of 
burn, blueberries were collected two years after pre- 
scribed fire from the ten burned over plots and from 
three control plots. Controls consisted of three clear- 
cut 35 m X 70 m plots adjacent to the burned-over 
plots. Blueberries were not collected in subsequent 
years because of logistical reasons limiting access to 
the research plots. 

For this study, three 3 m x 3 m quadrats were ran- 
domly established in each of the 35 m x 70 m plots. 
The perimeter of each of the 3 m x 3 m quadrats was 
delineated with a string tied 40 cm above ground and 
all berries within the 3 m x 3 m quadrats were picked, 
advancing from the edge of the plot inwards. Ripe, 
over-ripe, green, and deformed blueberries were in- 
cluded in the harvest. All plots were harvested on 13 
July and 14 July 1993. The harvested berries were kept 
in plastic coolers in the shade until the end of the col- 
lection day and their fresh weight determined in the 
laboratory the same evening. The bags were then placed 
in drying ovens at 50°C for 5 days before weighing. 
Because the moisture content of blueberries (83.1 + 
0.8 %) was comparable among all plots (P = 0.22) 
and consistent with published data (Usui et al. 1994), 
the results were expressed in terms of fresh weight only. 
As well, no attempt was made to distinguish between 
V. myrtilloides and V. angustifolium as the frequency 
of these species (1:15- 1:30, myrtilloides: angustifoli- 
um) was not significantly different among treatments 
(S051): 

To compare the effect of fire intensity and depth of 
burn on the vegetative abundance of V. myrtilloides and 
V. angustifolium, the number of stems of both species 
was determined two years after prescribed fire from 
the ten burned over plots and three control plots. Once 
the berries had been removed from the 3 m x 3 m 


DUCHESNE and WETZEL: EFFECT OF FIRE INTENSITY AND DEPTH OF BURN ON BLUEBERRY 


197 


quadrats, the number of stems was recorded for the 
entire quadrat and expressed as stems/m/?. 

As well, the number of blueberries/stem was com- 
pared among the prescribed burns and the control. 
For this, 100 stems were selected randomly outside 
the 3 m x 3 m quadrats and the number of blueberries 
determined for each stem. This protocol was repeated 
in each of the ten prescribed burn plots as well as each 
of the three control plots. 


Statistical analyses 

Analysis of variance (ANOVA) was conducted to 
compare blueberry production, stem density and blue- 
berry abundance/stem among the prescribed burns 
and the controls (Systat 1997). For statistical analyses 
the three control plots were pooled as one treatment 
whereas each of the ten burned over plots was treated 
as a separate treatment (McAlpine 1995). Means were 
compared using the Bonferroni procedure (Systat 
1997). Pearson’s multiple correlation analysis was con- 
ducted to determine the relationship between frontal 
fire intensity, depth of burn, berry production and the 
number of blueberries/stem (Systat 1997). 


Results 

Blueberry production (g/m?) differed significantly 
(P < 0.001) among treatments. The greatest blueberry 
production was observed in prescribed fires of 597 
and 1268 kW/m whereas there were no significant 
differences among blueberry production of the other 
treatments, including controls (Table 1). Likewise stem 
density/ha differed among treatments (P < 0.001) and 
was greatest in the fires of 597 and 1268 kW/m. There 
was no difference in the number of blueberries/plant 
among the other treatments including the controls 
(P = 0.056). 

Pearson’s multiple correlation analysis showed that 
fire intensity correlated (P < 0.05) with blueberry pro- 
duction, and stem density (Table 2). Depth of burn 
correlated (P < 0.01) with blueberry production, stem 
density and blueberries/plant. Depth of burn showed 
better correlations with blueberry production, number 
of blueberries/plant and stem density than fire inten- 
sity (Table 2). 


Discussion 

Several studies have demonstrated that the response 
of ecosystems varies greatly with frontal fire intensity 
and depth of burn. Fire intensity has been shown to 
correlate with P. banksiana regeneration (Weber et al. 
1987) and post-fire abundance of P. aquilinum (Tellier 
et al. 1995). Low fire intensities stimulated Corylus 
cornuta sprouting and height growth of Rubus ideaus 
(Johnston and Woodard 1985). Biomass nutrient reten- 
tion (Duchesne and Tellier 1997) and soil seed bank and 
competing vegetation dynamics along with P. resinosa 
and P. strobus seedling performance are also affected 
by prescribed burning intensity (Tellier et al. 1995, 
1996; Whittle et al. 1997). Soil microbial activity and 


198 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE 2. Pearson’s multiple correlation analysis among fire intensity, depth of burn, blueberry biomass, number of blueberries/ 


stem, and stem density. 


Intensity Blueberries/ha 
Intensity 1.000 
Blueberries/ha -0.507* 1.000 
Depth of burn 0.517% -0.683** 
Blueberries/stem -0.461 0.803** 
Stem density -0.504* 0.766** 


* significant at P < 0.05 
** significant at P < 0.01 


diversity were also affected by fire intensity (Duchesne 
and Wetzel 2000; Staddon et al. 1998a, 1998b). 

Fire has been used for hundreds of years in blue- 
berry management by Indigenous people (Chapeskie 
2001) and by farmers (Badcock 1958; Blatt et al. 
1989), mostly using spring fires. This investigation 
presents the first comprehensive data on the effect of 
fire intensity and depth of burn on natural blueberry 
production, which are indicated as critical factors to 
consider in blueberry management. Indeed, the current 
findings show that fire intensity must be kept low in 
order to increase blueberry productivity using pre- 
scribed fire. As well, care must also be given to reduce 
depth of burn, which has a greater impact on blueberry 
production than fire intensity (Table 2). Whereas fire 
intensity is a measure of the caloric energy liberated by 
the fire front (Alexander 1982), depth of burn integrates 
the interaction between soil moisture conditions and 
fire intensity (Miller 1977) and is more closely linked 
to the response of blueberries to fire than fire intensity. 

Since blueberry shoots arise from shallow rhizomes 
and portions of the above ground stem not killed by 
the fire (Flinn and Wein 1977; Martin 1955: Minore 
1975), deep heat penetration in the humus layers is 
likely to reduce stem production. Similar conclusions 
have been reached in Ponderosa Pine (Pinus ponder- 
osa) forests where depth of burn influenced greatly 
understory species regeneration (Armour et al. 1984) 
which is consistent with the concept that depth of 
buried propagules, together with heat penetration, is 
critical in post-fire survival (reviewed by Whittle et al. 
1997). In turn, the differences observed in this inves- 
tigation in post-fire blueberry stem densities, and the 
negative correlation between stem density and depth 
of burn may be ascribed to the deleterious effect of 
fire on the underground rhizomes as was demonstrated 
on Blue Huckleberry (Vaccinium globulare) (Miller 
L977): 

In practice, blueberry production can be maximized 
by burning in the spring of every second year, presum- 
ably to maintain a high ratio of flower buds to leaf buds 
(Hall et al. 1972). In this investigation, the number of 
blueberries/plant, which was hypothesized to be an 


Depth of burn Blueberries/stem Stem density 
1.000 
=O ei 1.000 
-0.724** 0.471 1.000 


estimate of the flower bud to leaf bud ratio, was not 
significantly different among treatments but inversely 
correlated with fire intensity. These results cannot be 
explained by conventional thinking regarding flower 
bud formation (Hall et al. 1972), as the number of blue- 
berries/plant correlated with depth of burn. However, 
Smith and Hilton (1971) speculated that Lowbush 
Blueberry performance after fire pruning might result 
from the stimulative effects of the nutrients in ash 
deposited on the surface soil during burning. Further 
investigation is needed to assert the relevance of this 
hypothesis in blueberry flower biology. 

Although this investigation shows that low intensity 
fires have a positive influence on blueberry production, 
natural blueberry ecosystems support a variety of wild- 
fire characteristics (Moola et al. 1998; Usui et al. 1995). 
Hence the continuous effect of repeated fires should be 
investigated on the long term productivity and biologi- 
cal conservation of such ecosystems. In particular, it 
will be important to assess the impact of burning every 
second year, which is the recommended practice, on 
soil nutrients (Smith and Hilton 1971). Future work 
should also be conducted to investigate co-manage- 
ment of blueberries with timber values (Duchesne et 
al. 2001). 


Documents Cited (marked * in text) 
Agriculture and Agri-Food Canada. 2002. Canadian fruit 
situation and trends. http://www.agr.gc.ca/misb/hort. 


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THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Whittle, C. A., L. C. Duchesne, and T. Needham. 1997. 
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Received 12 June 2002 
Accepted 24 September 2004 


Spring Dispersal Patterns of Red-winged Blackbirds, A gelaius 
phoeniceus, Staging in Eastern South Dakota 


H. J. Homan!, G. M. Linz!, R. M. ENGEMAN?, and L. B. PENRyY! 


'U.S. Department of Agriculture, National Wildlife Research Center, 2110 Miriam Circle, Bismarck, North Dakota 58501- 
2502 USA 

2 U.S. Department of Agriculture, National Wildlife Research Center, 4101 LaPorte Avenue, Fort Collins, Colorado 80521 - 
2154 USA 


Homan, H. J., G. M. Linz, R. M. Engeman, and L. B. Penry. 2004. Spring dispersal patterns of Red-winged Blackbirds, 
Agelaius phoeniceus, staging in eastern South Dakota. Canadian Field-Naturalist 118(2): 201-209. 


Red-winged Blackbirds (Agelaius phoeniceus) are very abundant summer residents throughout the Prairie Pothole Region 
of central North America. In late summer they assemble in post-breeding flocks that cause significant amounts of agricul- 
tural damage, particularly in sunflower fields near natal sites. In April 2001, we aerially color-marked ~370 000 Red-winged 
Blackbirds near Badger, South Dakota (44°48'N, 97°21'W), to determine if migrants staging here were summer residents in 
sunflower production areas ~350 km to the northwest. We measured patterns of migratory dispersal by collecting birds in 54 
randomly selected blocks in the northcentral U.S. and the Prairie Provinces of Canada. The marked specimens (n = 33) were 
categorized into three polygons based on analyses of banding and re-sighting data and proximity to concentrated sunflower 
production. We estimated that 82% of the migrants that had staged in eastern South Dakota resided. within or on the 
periphery of the sunflower growing area. These birds probably stay near their breeding territories until at least late August 
and cause early damage to sunflower, which comprises the majority of damage. Resident birds in Alberta and most of 
Saskatchewan (18%) might arrive too late in the damage season to impact the sunflower crop significantly. 


Key Words: Agelaius phoeniceus, Red-winged Blackbird, breeding range, color-marking, dispersal patterns, northern Great 


Plains, spring migration, sunflower damage. 


Several million hectares of wetlands have been 
drained in the Prairie Pothole Region (Dahl 1990*), but 
high densities of semipermanent wetlands remain in 
North Dakota (0.88/km?, Reynolds et al. 1997*). These 
are preferred nesting habitat of Red-winged Black- 
birds (Agelaius phoeniceus) and make North Dakota 
the center of abundance for this species. During the 
mid-1990s, the breeding population in North Dakota 
started gaining rapidly in size. Red-winged Blackbird 
indices from the North American Breeding Bird Sur- 
vey (BBS) are at historical highs for the state (Figure 
1). From 1994-2002 North Dakota’s average BBS index 
of all routes in the Prairie Pothole Region ranged 
from 199 to 250 Red-winged Blackbirds per route. 
North Dakota led all other U.S. states and Canadian 
provinces during this period in annual indices of Red- 
winged Blackbirds (Sauer et al. 2004*). Conflicts bet- 
ween wildlife and agriculture often occur because of 
local overabundance (Garrott et al. 1993; Conover 
2002), and the growth in numbers of Red-winged 
Blackbirds has exacerbated a long-term struggle bet- 
ween sunflower producers and blackbirds (Linz and 
Homan 1998). Oilseed sunflower is a preferred food 
of Red-winged Blackbirds (Linz et al. 1984; Homan 
et al. 1994). It is also an important economic crop in 
North Dakota. In 2001, 75% of total oilseed sunflower 
production in North America came from North Dakota 
(National Agricultural Statistics Service 2004*). With 
increased depredation pressure on the crop caused by 
population expansion of Red-winged Blackbirds, many 


growers probably dropped sunflowers from their field 
rotations (Lamey and Luecke 1994). Since 1995, the 
number of sunflower fields has steeply declined in cen- 
tral and southeastern North Dakota, areas renowned for 
their high levels of sunflower production (National 
Agricultural Statistics Service 2004*). 

Statewide field damage surveys have that blackbird 
damage to sunflower ranges from $4 to 11 million 
(U.S. dollars) in the Prairie Pothole Region (Hothem 
et al. 1988). Based on current population estimates and 
bioenergetic models, Red-winged Blackbirds cause 
$3 million of the $5.5 million lost yearly to blackbirds 
in the core area of sunflower production (Peer et al. 
2003). Most of the damage occurs between mid-August 
and early September, when the calorie content of im- 
mature sunflower achenes is low and the birds must 
eat more to reach satiation (Cummings et al. 1989; Con- 
ner and Hall 1997). During this early damage period, 
when >75% of total damage occurs, Red-winged 
Blackbirds account for 80% of the blackbird species 
observed in sunflower fields (Cummings et al. 1989). 
This period is before en masse blackbird migration, and 
the majority of these birds are of local origin. Most will 
remain within 200 km of their natal sites until molt 
completion (or near completion) in late August and 
early September (Dolbeer 1978; Besser et al. 1983; 
Linz et al. 1983). The damage can be quite serious in 
central and southeast North Dakota and northeast South 
Dakota, areas of concentrated production of sunflow- 
ers (National Agricultural Statistics Service 2004*) and 


201 


202 


plentiful wetlands to attract nesting Red-winged Black- 
birds (~22 breeding pairs/km7, Linz et al. 2002). By 
late summer, the Red-winged Blackbird population 
responsible for most of the sunflower damage in the 
Prairie Pothole Region is about 39 (+8.8) million (Peer 
et al. 2003). 

Reducing excessive field damage (e.g., >10% per 
field) has proven an enormous challenge. The U.S. 
Department of Agriculture (USDA) uses an integrated 
management approach of lethal and nonlethal methods. 
Most of the available methods are used infrequently 
because of unmanageable logistics, little or no efficacy, 
or low cost:benefit ratios (Linz and Hanzel 1997). 
Mechanical frightening agents (e.g., scarecrows, pyro- 
technics, and propane cannons) help alleviate light 
predation pressure, but only work effectively if the dur- 
ation of the damage period is less than the habituation 
period (Cummings et al. 1986). Unfortunately, black- 
birds habituate quickly to frightening agents, particu- 
larly if the crop is a preferred food in an area of limited 
alternate foods (Ward 1979). The taste repellent, methyl 
anthranilate, is also used by producers, but the con- 
centration in the commercial product (BirdShield™) 
is below the known repellency threshold, despite the 
formulation’s high cost (Werner et al. 2005). Experi- 
ments with avicide were tried in sunflower fields with 
ongoing damage. Avicide-treated rice (DRC-1339) 
was placed in small plots opened in the fields. This 
approach failed because (1) blackbirds foraged almost 
exclusively on the standing heads of sunflower and 
would not go to the ground to forage in the plots, (2) 
precipitation necessitated numerous rebaitings, and 
(3) nontarget avian risks increased substantially over 
the baiting period because of migration (Schaaf 2003; 
Linz et al. 2000). Two nonlethal methods showing 
some promise are reduction of dense stands of cattail 
(Typha spp) in wetlands near sunflower (Linz et al. 
2004a) and strategic placement of lure crops to attract 
foraging blackbirds away from sunflower fields (Linz 
et al. 2004). Cattail management has been a USDA 
program for over 10 years. The program is small com- 
pared to existing coverage of cattail (Ralston et al. 
2004); and it also faces year-to-year logistical chal- 
lenges because of instability in roost locations across 
years combined with cattail’s ability to rapidly invade, 
colonize, or re-colonize wetlands (Homan et al. 2003). 
Research on efficacy of lure crops is just beginning 
and time will be needed to assess this method. 

Sometimes population management may be the most 
realistic and effective method for reducing persistent 
damage caused by overabundant wildlife populations 
(Garrott et al. 1993). The definition of overabundance 
is subjective and based upon one’s interests, likes, and 
dislikes; and therefore is controversial. It has been 
defined generally as that point of abundance where all 
positive values of the species have been overwhelmed 
by negative values created by the sheer numbers of indi- 
viduals (Conover 2002). The definition implies both a 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


spatial and temporal component, as populations are 
seldom ubiquitous or always aggregated. The defini- 
tion is fitting, at least from an agricultural perspective, 
in the sunflower growing areas of North Dakota and 
northeastern South Dakota. The USDA has drafted an 
Environmental Impact Statement (EIS) that addresses 
the conflict between sunflower producers and black- 
birds in North Dakota, South Dakota, and Minnesota 
(Federal Register 2001*). Several management options 
were included in the draft EIS. The research described 
herein addresses one of the options being considered: 
population management of spring migrating Red- 
winged Blackbirds in northeastern South Dakota. This 
part of South Dakota is one of the most northward sites 
where the population is still grouped together enough 
to make population management a feasible alternative. 

Because blackbird damage to sunflower is a local- 
ized phenomenon occurring within the broader region 
of the northern Great Plains (Peer et al. 2003), it is nec- 
essary to identify specific origins of local populations 
before implementing any strategy of population man- 
agement. Although leg-banding has provided valu- 
able insights into migratory timing and dispersal of 
Red-winged Blackbirds (Dolbeer 1978), this method 
draws its samples randomly from regional populations. 
Moreover, it is labor intensive in both its capturing 
and ringing methods; these drawbacks combined with 
natural annual mortality of Red-winged Blackbirds 
cause data to be parsed over several years because of 
extremely low rates of band returns (<0.5%, Besser et 
al. 1983; Gammell et al. 1986; Stehn 1989). An alterna- 
tive method of tracking movements, aerial mass color- 
marking, provides a means for rapid accumulation of 
data on movements of local populations within a single 
season framework. Recovery rates from mass color- 
marking projects depend on the number marked com- 
pared to the size of the study population (Johns et al. 
1989): however, thousands of birds can be marked in 
a single application often yielding a rate of return 10 
to 20 times that of leg-banding. Aerial mass color- 
marking has been used to track spring movements of 
Red-winged Blackbirds from northeastern Missouri 
and eastern South Dakota (Knittle et al. 1987; Knittle 
et al. 1996), short late-summer movements of Red- 
winged Blackbirds in central North Dakota (Linz et 
al. 1991), and movements of Red-winged Blackbirds 
between winter roosts in the southcentral U.S. (Harsch 
1995*). The marking formulation is environmentally 
safe at the concentrations used (Bills and Knittle 1986; 
Knittle and Johns 1986). 

In April 2001, we used aerial mass-marking to iden- 
tify migrating Red-winged Blackbirds staging in large 
cattail-dominated wetlands in northeastern South Da- 
kota. Millions of spring migrating blackbirds annu- 
ally stage at this traditional stopover site between late 
March and April (Sawin 1999). Our objective was to 
assess migratory dispersal to breeding territories north 
of the marking site. Previous research has shown that 


2004 HoMaAn, LINZ, ENGEMAN, and PENRY: DISPERSAL PATTERNS OF BLACKBIRDS 203 

250 

200 

North Dakota 
L 
5 “, 
a 150 ree % 
= Fi 
g NL * Saskatchewan 
7) oe ‘ 
— « Le 
= 5 a _Manitoba 
8 100 rid PO eee ee a ims 
” i 
a Fd 
a ” 
© Alberta 
= 50 
0 
< 2 2 2 ND 2 £ 
Year 


Ficure 1. Three-year rolling averages of indices from the North American Breeding Bird Survey from 1967 to 2003. The in- 
dices are from routes in the Drift Prairie, Glaciated Missouri Plateau, and Aspen Parklands, which are the three major 
physiographic regions in the Prairie Pothole Region used most frequently by Red-winged Blackbirds. 


color-marked male Red-winged Blackbirds staging 
in eastern South Dakota during spring were in part 
destined for breeding grounds in or near concentrated 
sunflower production; however, these data were col- 
lected over 15 years ago (Knittle et al. 1987; Knittle 
et al. 1996). Population management programs, when 
used at sites far from the locality where actual damage 
occurs, must be based on a thorough and contempo- 
rary knowledge of movements of individual popula- 
tions (Dolbeer 1978). We believed it necessary to repli- 
cate the marking studies because changes may have 
occurred in migratory patterns. Additionally, no data 
exist on dispersal movements of females, and these 
data would increase our understanding of the popula- 
tion’s migration patterns. 


Methods 
Marking 

During March 2001, we monitored cattail-dominat- 
ed wetlands in six counties in eastern South Dakota 
for arrival of migratory flocks of blackbirds. Three wet- 
lands held migratory roosts large enough (>10 000 
birds) to warrant aerial mass-marking. All three roosts 
were within a 32-km radius of Badger, South Dakota 
(44°48'N, 97°21'W). Roost size was estimated by 


counting blackbirds as they entered the roost on the 
evening of the spray (Meanley 1965; Arbib 1972). Spe- 
cies compositions and sex compositions were estimat- 
ed by randomly selecting and identifying groups of 
individual birds as they entered (Dolbeer et al. 1978). 
A fixed-wing agricultural spray plane applied the 
marker near dark at an altitude of 20—30 m (Knittle et 
al. 1987). The following morning we collected birds 
as they departed the sprayed roost and later examined 
them for fluorescent marks using a long-wave (360 nm) 
ultraviolet light. We calculated the number of marked 
male and female Red-winged Blackbirds in the roost 
by multiplying the proportion of marked birds in the 
sample by the estimated number of males and females 
using the roost on the night of the spray. 

The marker formulation was a 416-L solution con- 
sisting of 208 L Carboset® (an acrylic adhesive; Nov- 
eon™ Incorporated, Cleveland, Ohio), 102 L food- 
grade propylene glycol, 7.7 kg fluorescent organic pig- 
ment (DayGlo® Color Corporation, Cleveland, Ohio), 
1 L each of a defoamer and surfactant, and 106 L 
water (Knittle et al. 1996). We used CP® nozzles set at 
5 (400 micron droplets). Spray volume was 28L/ha 
delivered at 1.8 kg/cm’. In spray form the solution 
dries in 2-3 minutes adhering to the feather surface. 


204 


Knittle and Johns (1986) found that 5% of marked birds 
lost their marks within six weeks; however, particles 
lodged in the barbules can remain until molt (Knittle 
et al. 1996; Otis et al. 1986). 


Sampling and Collections 

We grouped the study area into 140 blocks, each 
consisting of four 1°-latitude x 1°-longitude units. Two 
units were randomly selected from each block. To en- 
sure adequate sample sizes per effort, only those units 
within regions that averaged 230 Red-winged Black- 
birds per BBS route were selected (Sauer et al. 2004); 
37 units in Canada and 19 units in the U.S. were chos- 
en in this manner. No units south of the 1° x 1° unit 
containing the marking sites were sampled. The sam- 
ple units covered an extensive area (>1.0 x 10° km?) 
bounded by 44—57° N latitude and 95—119° W longi- 
tude (Figure 2). Southwestern North Dakota, central 
Minnesota, central and western South Dakota, and 
southeastern Montana were excluded because BBS 
counts were <30 (Sauer et al. 2004). 

We assigned the 1° x 1° units to one of three disper- 
sal polygons: core, peripheral, and outside-peripheral 
(Figure 2). The shape and area of the polygons were 
based on an analysis of banding and re-sighting data, 
physiography, and proximity to the area of concen- 
trated sunflower production (Stehn 1989). The core 
polygon encompassed the breeding range of birds hav- 
ing a high probability of causing early damage to sun- 
flowers. The peripheral polygon contained the sur- 
rounding breeding range of birds that were capable 
(with only short pre-migratory movements) of entering 
the core area during the early damage period (mid- 
August — early September) (Dolbeer 1978; Gammell 
et al. 1986). The outside-peripheral polygon contained 
the birds farthest removed from the core area and en- 
compassed the rest of the study area not covered by 
the core and peripheral polygons. The 1° x 1° units 
were placed in core and peripheral polygon categories 
if any of the unit’s area intercepted the polygon’s bound- 
ary. We used a GIS to place the polygons on a geo- 
referenced grid of the 1° x 1° units. 

We used shotguns (12-gauge with either #6 or #7/ 
steel shot) to collect male and female Red-winged 
Blackbirds on breeding territories during June in the 
U.S. and June and early July in Canada. The wings 
were removed from the specimens and placed in an 
envelope with the GPS coordinates of the collection 
site. The wings were later viewed for fluorescent pig- 
ments using a 360-nm ultraviolet lamp under 1.75 mag- 
nification. The markings on tagged birds are usually 
subtle and invisible to the naked eye under natural 
light. To eliminate false positives, only splash marks 
and individual particles firmly attached to the feathers 
were considered conclusive evidence of a valid mark 
(Knittle et al. 1987). Blood samples were removed on- 
site from randomly selected specimens and used for 
microsatellite genetic analysis in a separate study 
(Williams et al. 2004). The wings were frozen and shall 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


be kept for future research on primary feather chem- 
istry and its relationship to geographic origins (Ed- 
wards and Smith 1984). 

The data were analyzed using descriptive statistics, 
likelihood ratio chi-square analyses (G-tests), and non- 
parametric tests. We used G-tests to assess differences 
between (1) the proportion of marked birds (sexes com- 
bined) in the sample with the expected proportion of 
marked birds in the study area; (2) the proportions of 
marked males and females in the sample with the ex- 
pected proportions of marked males and females in the 
study area; and (3) the proportions of marked birds 
within core, peripheral, and outside-peripheral poly- 
gons. The first two tests used extrinsic expected fre- 
quencies based on estimates of marking efficiency at 
the staging site and population size in the study area 
during the breeding season. The third comparison used 
intrinsic frequencies generated exclusively from the 
collections (Sokal and Rohlf 1981). Population size 
in the study area was estimated using a combination 
of stable age structure analysis from banding data and 
linear regression analysis of BBS indices and inde- 
pendently determined density estimates (Stehn 1989). 
The resultant regression equation was Y = 0.3X, where 
X was the average BBS count within each ecological 
stratum in the study area and Y was the population den- 
sity per km? (see Aldrich 1963; Robbins et al. 1986). 
We used the Wilcoxon Rank Sum test to compare 
ranked differences between males and females in dis- 
tances traveled from the marking site to the collecting 
sites. Distance measurements were made using a GIS. 


Results 

We applied fluorescent marker seven times, tagging 
an estimated 235 500 male and 131 100 female Red- 
winged Blackbirds over the period 1-23 April. This 
period covered the Red-winged Blackbird migration 
through eastern South Dakota in 2001. We collected 
2398 males and 2060 females on breeding territories, 
of which 33 (0.74%) were marked (Table 1). The 
overall recovery rate of 0.74% was similar to the 
expected recovery rate of 0.96% (G = 1.1, P = 0.30), 
based on total number of birds marked in April and the 
estimated population size of breeding Red-winged 
Blackbirds in the sampled collection area (38.1 x 10°, 
from Stehn 1989) which included the population in 
the outside-peripheral polygon. The ratio of marked 
males : females in the sample (f = 23 : 10) was near 
the expected ratio of 22 : 11 (G = 0.07, P = 0.80). 
Proportions of marked birds collected in the three 
polygons differed (G = 6.44, P = 0.04), with the per- 
centages of marked birds in the core polygon (1.03%) 
and peripheral polygon (0.93%) nearly three times 
greater than the percentage collected in the outside- 
peripheral polygon (0.35%) (Table 1). The outside- 
peripheral polygon contributed 69% of the total G- 
value. Of the 33 marked recoveries, 15 (46%) were 
collected in the core polygon and 12 (36%) were 
collected in the peripheral polygon. 


2004 Homan, LINZ, ENGEMAN, and PENRY: DISPERSAL PATTERNS OF BLACKBIRDS 205 


Outside-peripheral 
polygon 


Peripheral polygon 


Core polygon 


Axel 
study block 


© Female N 
A Male A 


3x Marking site 300 0 300 Kilometers 


a okt 


FiGurRE 2. Locations of 33 marked Red-winged Blackbirds (in relation to core, peripheral- and outside-peripheral polygons) 


after dispersing from a spring staging site in eastern South Dakota in April 2001. 


TABLE 1. Color-marked male (©’) and female (Q) Red-winged Blackbirds collected during June and July 2001 in core, 
peripheral, and outside-peripheral polygons after migratory dispersal from a staging site in eastern South Dakota during 


April 2001. 

Number Number Number Percent 

units? Collected Marked Marked 

Polygon ‘os O Total ‘oi fe) Total ol Q All 
Core 14 793 668 1 461 9 6 15 eit} 0.90 1.03 
Peripheral 15 681 605 1 286 10 2 12 1.47 0.33 0.93 
Outside 27 924 787 Lala 4 D 6 0.43 0.25 0.35 
Total 56 2 398 2 060 4 458 2B 10 33 0.96 0.48 0.74 


4No 


birds were collected in two units 


206 


No marked birds were collected north of 53° N. 
Recovery distances were similar between sexes (T = 
0.18, P = 0.86), with the median location 519 km 
(range: 20-1353) from the marking sites (48°23'N, 
100°27'W). The median azimuth from the marking 
sites was 325°. The median distance and direction of 
marked females placed them on the northwestern side 
of the core polygon (48°23'N, 100°27'W; whereas, 
males were 70 km due west (48°30'N, 101°23'W), in 
the peripheral polygon. For all recoveries combined, 
the median location was 50 km northeast of Minot, 
North Dakota (48°14'N, 101°18'W). 


Discussion 

The seven sprays were evenly distributed through- 
out the migratory period and probably yielded an 
accurate depiction of the migratory distribution pat- 
terns of Red-winged Blackbirds. Similar to the results 
of previous banding and color-marking experiments 
in eastern South Dakota, our data showed that most 
birds followed the Prairie Pothole Region as they 
traveled northwest to their breeding territories (Besser 
et al. 1983; Knittle et al. 1987; Knittle et al. 1996). 
Although Red-winged Blackbirds are not confined to 
breeding in this region, they are found here in their 
highest densities. A statewide population census of 
breeding birds in North Dakota showed that the Prairie 
Pothole Region (comprised of several physiographic 
regions, but mainly the Drift Prairie and Glaciated 
Missouri Plateau) contained 70% of the state’s Red- 
winged Blackbird population (Stewart and Kantrud 
1972). 

Unlike parameters such as median distance and 
direction, both of which can be influenced by differ- 
ences in breeding densities over the study area, the pro- 
portions of marked birds within the three polygons were 
distribution measures mathematically independent of 
density. It was thus possible to consider our results as 
reflections of behavioral phenomena, such as fidelity 
to migratory pathways and philopatry. The marked 
population was apparently not distributing itself ran- 
domly in the study area. The low proportion of marked 
birds in the outside-peripheral polygon suggested that 
eastern South Dakota may be used mainly by migrants 
already near their final destination, and as a result, 
traveling relatively short distances after departure from 
the staging site. We speculate that other staging sites 
might exist (perhaps much farther to the west), and 
these should have birds destined for breeding territories 
in Montana, southern Alberta, and Saskatchewan. 
Marking of large roosts farther west (for example, a 
large blackbird roost exists near Great Bend, Kansas 
[38° 22'N, 98° 49'W]) may demonstrate that western 
migratory pathways exist distinct from the one used by 
blackbirds in eastern South Dakota. Certainly, the very 
low proportion of marked birds in the outside-periph- 
eral polygon, compared to the other two dispersal 
polygons, suggests a different population of birds. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


There were two other marking studies done in the 
general vicinity of our marking site. These were done 
at Lake Thompson, South Dakota (44°23'N, 97°33'W), 
approximately 30 km to the southwest. Both were 
similar in design, scope, and sampling effort to our 
study, except that only males were collected. One study 
was conducted in 1983 (Knittle et al. 1987), the other 
in 1985 (Knittle et al. 1996). Even though geopolitical 
boundaries were used to analyze returns instead of dis- 
persal polygons, comparisons of results among and 
between studies can still be made. Results were similar 
between our study and the 1983 study in two impor- 
tant facets: (1) the recovery rate of marked birds in 
Canada was substantially lower than the U.S. recov- 
ery rate (4.4 vs. 7.0%, Knittle et al. 1987), and (2) 
proportions of marked birds collected in western North 
Dakota, central North Dakota, and northeastern South 
Dakota were greater than other regions. By contrast, 
results diverged in the 1985 study, both from our results 
and from those from the 1983 study. The expected 
rate of recovery in the 1985 study was much less than 
the observed recovery rate (5.2 vs. 14.6%, Knittle et 
al. 1996). To obtain a 14.6% recovery rate, the male 
population would have had to have been 6.7 x 10°, a 
number 65% lower than what was estimated in the 
study area during that time (19.0 x 10°, Stehn 1989). 
Saskatchewan’s Red-winged Blackbird population 
had indeed been in steep decline for nearly a decade, 
as evidenced by BBS indices (Figure 1); however, 
trends for Alberta, Manitoba, and North Dakota were 
either stable or increasing. It is unlikely for such a 
conspicuous species that a 65% decline would be 
missed in the BBS surveys. Of course, the number of 
birds marked in 1985 could have been underestimated 
by this magnitude, but an even more anomalous result 
in the 1985 study argues against this; the percentage 
of marked birds in Canada was greater than the per- 
centage marked in the Prairie Pothole Region below 
49° N (17 vs. 16%). Usually, recovery rates are inverse- 
ly related to the distance traveled from a marking site 
because unmarked populations arriving by other routes 
from other areas dilute the proportion of marked birds. 
Although it is conceivable that the Red-winged Black- 
bird population marked at Lake Thompson in 1985 
migrated beyond traditional nesting areas and immi- 
grated into Canada, perhaps seeking better wetland 
conditions, band recoveries collected over several dec- 
ades have shown that Red-winged Blackbirds usually 
return to within 100 km of their natal sites (Nero 
1956; Dolbeer 1978). If Red-winged Blackbirds were 
not philopatric, but nomadic or irruptive, we contend 
that large fluctuations in BBS counts would occur 
among physiographic regions comprising the Prairie 
Pothole Region. No significant correlations existed in 
yearly changes in BBS indices between North Dakota 
and the three Prairie Provinces within our study area, 
and perusal of average BBS indices over a 36-year 
period does not seem to indicate irregular migratory 


aegnnce 


behavior (Figure 1). The extraordinarily high overall 
recovery rate of marked birds in the 1985 study may 
have resulted from improper identification of marks 
(i.e., false positives) caused by environmental contam- 
ination with fluorescence pigments used for identifying 
insecticide formulations and fertilizers (Knittle et al. 
1987). We believe that our data from 2001 and the data 
from the 1983 study have provided the most accurate 
analyses of dispersal patterns of migrating Red-winged 
Blackbirds in eastern South Dakota. 


Summary and Conclusions 

In landscapes with numerous wetlands, blackbird 
damage to sunflowers averages 25% per field. One- 
quarter of the fields receive >10% damage (Linz et al. 
2000). At the 10% damage level, agricultural producers 
start dropping sunflower from their crop rotations and 
replacing it with other oil crops, such as soybean and 
canola (North Dakota Agricultural Statistics 1990*; 
Lamey and Luecke 1994). There is no evidence that 
the rapid population growth of Red-winged Blackbirds 
has directly affected other bird species; however, the 
potential for indirect effects exists. Sunflower fields 
were used by 49 nonblackbird species, representing all 
the major feeding guilds, during the late-summer and 
early fall migrations (Schaaf 2003). Sunflower fields 
have several characteristics that could make them excel- 
lent migratory stopover sites for rest, cover, and pro- 
curement of energy reserves (Hutto 1998; Petit 2000). 
First, they provide large blocks of sturdy herbaceous 
cover with a dense canopy and rank understory in a 
landscape that by mid-August is mostly barren north of 
46°N because of the harvest of small grains (Schaaf 
2003). Second, sunflower fields provide a diverse sel- 
ection of foods, including seeds of weedy plants in the 
furrows and numerous invertebrates in and around 
the fields (Schulz 1978; Charlet et al. 1997; Schaaf 
2003). This attracts avian granivores and insectivores 
(and of course, the carnivores). Soybean and canola 
fields, which often replace sunflower, have neither the 
beneficial vegetative structure nor the energy and are 
a less productive habitat for birds (Linz et al. 2004d). 
Moreover, these crops are “clean cropped,” unlike sun- 
flower, which is often not plowed until May of the fol- 
lowing year. Sunflower stubble was used by 33 species 
of migrating birds during early spring in North Dakota 
(Galle et al. 2004). It was used more frequently 
during spring migration than fields of soybean, small 
grain, corn, and sorghum. Thus, loss of sunflower fields 
in the Prairie Pothole Region could have unforeseen 
ecological impacts on migrating nonblackbird species. 

We estimate that 82% of the Red-winged Blackbirds 
that stage in eastern South Dakota during spring migra- 
tion reside either within or on the periphery of the sun- 
flower growing area. If population management of 
the spring breeding population in the northern Great 
Plains is deemed the best of the EIS alternatives, we 
suggest that the most efficacious strategy would be to 


HoMAN, LINZ, ENGEMAN, and PENRY: DISPERSAL PATTERNS OF BLACKBIRDS 


207 


manage the population in eastern South Dakota. Lastly, 
it may be possible to avoid impacting the Canadian 
migrants, which comprised 18% of the birds staging in 
eastern South Dakota, by not applying management 
over the entire migration. Canadian Red-winged Black- 
birds should be among the last of the regional popula- 
tions of Red-winged Blackbirds to arrive at the stag- 
ing site (Weatherhead and Bider 1979). The arrival of 
Yellow-headed Blackbirds (Xanthocephalus xantho- 
cephalus), a nontarget blackbird species, usually signi- 
fies that the Red-winged Blackbird migration is nearing 
its end and may be a good indicator to cease manage- 
ment operations. 

We recommend future research on (1) genetic 
markers to identify local populations of Red-winged 
Blackbirds residing in Canada and the U.S.; (2) 
color-marking spring migrants in flyways farther west; 
(3) the relationship between timing of arrival at stag- 
ing sites and distance traveled to breeding territories; 
and (4) color-marking pre-migratory roosts [or other 
means of identification, such as genetics or mineral 
analyses of primary feathers] northwest of the core 
sunflower growing area prior to fall migration. 


Acknowledgments 

Personnel from North Dakota/South Dakota Wildlife 
Services and North Dakota State University helped 
collect the birds. We thank the following for their sup- 
port and cooperation on this project: the Departments 
of Agriculture of North Dakota and South Dakota; the 
U.S. Fish and Wildlife Service; Minnesota Department 
of Natural Resources; North Dakota Game and Fish 
Department; South Dakota Department of Game, Fish 
and Parks; South Dakota Department of Environment 
and Natural Resources; Montana Department of Fish, 
Wildlife and Parks; Canada Customs and Revenue 
Agency; Canada Department of Foreign Affairs and 
International Trade; the Environmental Protection 
Branch of Environment Canada; the Wildlife Branch 
of Manitoba Conservation; the Environment Branch 
of the Alberta Natural Resource Service; the Fish and 
Wildlife Branch of Saskatchewan Environment and 
Resource Management; the Manitoba Wildlife Animal 
Care Committee and Animal Care Committees within 
other provincial, federal, and state governments; the 
National Sunflower Association; and private landown- 
ers in the U.S. and Canada. Our research was conduct- 
ed under the auspices of Quality Assurance Study Pro- 
tocol QA-776, as amended; as such, safety guidelines, 
animal care, and scientific validity were reviewed, 
approved, and permitted by the National Wildlife 
Research Center, the U.S. Fish and Wildlife Service, 
the Environmental Protection Branch of Environment 
Canada, and the natural resource agencies in the states 
and provinces within the study area. Use of commercial 
products by the U.S. Government does not constitute 
an endorsement of these products. 


208 


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Received 30 July 2002 
Accepted 29 October 2004 


Changes in Loon (Gavia spp.) and Red-necked Grebe (Podiceps 
grisegena) Populations in the Lower Matanuska-Susitna Valley, Alaska 


TAMARA K. Mitis!* and BRAD A. ANDRES2* 


' Department of Biological Sciences, University of Alaska Anchorage, 3211 Providence Drive, Anchorage, Alaska 99508 
USA 

2 U. S. Fish and Wildlife Service, Nongame Migratory Bird Management, 1011 East Tudor Road, Anchorage, Alaska 99503 
USA 

> current address: U. S. Fish and Wildlife Service, Nongame Migratory Bird Management, 1011 East Tudor Road, Anchorage, 
Alaska 99503 USA 

* current address: Division of Migratory Bird Management, U. S. Fish and Wildlife Service, P.O. Box 25486, DFC-Parfet, 
Denver, Colorado 80225-0486 USA; corresponding author 


Mills, Tamara K., and Brad A. Andres. 2004. Changes in loon (Gavia spp.) and Red-necked Grebe (Podiceps grisegena) 
populations in the lower Matanuska-Susitna Valley, Alaska. Canadian Field-Naturalist 118(2): 210-214. 


More than two-thirds of the human population of Alaska resides in the south-central portion of the state, where its continued 
growth is likely to affect some wildlife populations negatively. To assess changes in waterbird populations in this region, we 
compared counts of Common Loons (Gavia immer), Pacific Loons (G. pacifica), and Red-necked Grebes (Podiceps 
grisegena) made on Matanuska-Susitina Valley lakes. In general, the number of lakes occupied by loon or grebe pairs 
decreased between 1987 and 1999. Decreases in the number of lakes occupied by Common Loons were less drastic in the 
northwest region of the study area than in the southeast region; human development is greater in the southeastern portion of 
our study area. Contrary to lake occupancy, the percentage of lakes that fledged Common Loon chicks remained stable between 
years. Because the human population is expected to continue to grow, proactive management of lake use and lakeshore 
development, coupled with monitoring of loon and grebe occupancy and productivity, is needed to ensure the persistence of 
these waterbird populations in the lower Matanuska-Susitna Valley. 


Key Words: Common Loon, Gavia immer, Pacific Loon, Gavia pacifica, Red-necked Grebe, Podiceps grisegena, breeding 


occupancy, productivity, south-central Alaska. 

The stability of many bird populations mainly de- 
pends on the stability of the environments they inhabit 
(Newton 1998). Nesting loons (Gavia spp.) and grebes 
(Aechmophorus spp., Podiceps spp.) are often nega- 
tively affected by habitat degradation and increased 
levels of human-related disturbances (Riske 1976; 
McIntyre 1978; Evers 2003). Common Loons (Gavia 
immer), Pacific Loons (G. pacifica), and Red-necked 
Grebes (Podiceps grisegena) breeding on lakes in the 
Matanuska-Susitna Valley (Mat-Su Valley), Alaska, are 
encountering many changes to their local environment. 
The area’s human population has tripled in the last two 
decades, and the current annual growth rate is 3.3% 
(McKibben and Nelson 1999*). Within the Mat-Su 
Valley, population growth has been greatest in the 
southeast, the area closest to Alaska’s largest city 
(Anchorage; McKibben and Nelson 1998*, 1999*),. 
The growing human population in the Mat-Su Valley 
has begun to encroach upon the freshwater habitat 
favored by breeding loons and grebes. Development of 
lake shorelines for houses and increased recreational 
use of lakes are suspected of having detrimental effects 
on loon and grebe populations in the Mat-Su Valley 
(Tankersley 1987*, Fair 1998*). 

Concern over the susceptibility of local loon popu- 
lations to anthropogenic pressures prompted the forma- 
tion of a volunteer program, the Alaska Loon Watch, 


in 1985. Under the direction of the Alaska Department 
of Fish and Game (ADF&G), volunteers collected data 
on loon and grebe presence and breeding activity on 
lakes throughout the Mat-Su Valley (see Fair 1998*). 
Using data collected through the Alaska Loon Watch 
program and our own observations, we assessed changes 
in lake occupancy, distribution, and productivity of loons 
and grebes breeding in the Mat-Su Valley between 
1987 and 1999. Based on the significant growth of the 
human population in the region, we predicted that lake 
occupancy and productivity by loons and grebes would 
have decreased over time. Furthermore, we predicted 
that the distribution of loons and grebes would shift 
spatially from southeast to northwest in response to 
corresponding changes in intensity of shoreline devel- 
opment and lake recreation. 


Study Area 

The Mat-Su Valley of south-central Alaska lies be- 
tween the Matanuska and Susitna rivers and borders 
upper Cook Inlet. Over 1000 lakes, ponds, and wet- 
lands of glacial origin occupy kettles in moraines and 
ice-stagnation complexes in this 500 km? region 
(Colazzi et al. 1986°). Lakes are classified as oligo- 
trophic or mesotrophic with pH levels conducive to 
support fish species preyed on by loons and grebes 
(Ruggles 1991). The elevation of the valley floor ranges 
from tide level to 400 m, but local relief of the area is 


210 


2004 


Uplands are dominated by White Spruce (Picea glauca), 
Paper Birch (Betula papyrifera), and Quaking Aspen 
(Populus tremuloides), Black Cottonwood (Populus 
trichocarpa) and willows (Salix spp.) are common 
along waterways and on alluvial plains. Maritime in- 
fluences moderate the climate in the Mat-Su Valley 
with average mid-summer temperatures of 14°C, annu- 
al snowfall of 150 cm, and annual precipitation of 
about 48 cm. We concentrated our survey effort with- 
in the lower Mat-Su Valley, an area bounded by the 


MILLS AND ANDRES: CHANGES IN LOON AND GREBE POPULATIONS IN ALASKA 


211 


communities of Palmer to the southeast, Willow to 
the northwest, Sutton to the east, and the Susitna 
River to the west (Figure 1). 


Methods 

We surveyed lakes in late May and late August of 
1999 to determine occupancy by loons and grebes 
and productivity of loons. Because of their smaller size 
and tendency to linger in aquatic vegetation, produc- 
tivity of Red-necked Grebes was not recorded. Of 


\ 


Palmer 
3) ¥ 


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ys > So & (a) 
= c Sans Eklutna 
oars 1 5 ys iG co N 
L b agg “ Sn kee 
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BN - « s * 4 €s V 
» ? 4 p as ae ya 
a .. OK 3 A Woe 
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a © 10 20 Kilometers 
Fee | 
: ne ea 


FiGurE 1. Location of the lower Matanuska-Susitna Valley, Alaska, where lake occupancy and productivity of loons and 
grebes were assessed, in the delineated area, in 1987 and 1999. 


D2 


219 lakes surveyed in 1987 by biologist Nancy 
Tankersley (ADF&G) and volunteers of the Alaska 
Loon Watch program, we determined that 139 lakes 
had adequate coverage in both years to allow a mean- 
ingful comparison. In 1999, observers scanned each 
lake with binoculars from several vantage points along 
the lakeshore to ensure complete lake coverage. Lakes 
were scored for the presence or absence of single loons 
or loon pairs during late May and for the presence or 
absence of loon chicks during late August. Field meth- 
ods were similar in 1987. A lake was considered oc- 
cupied if a loon pair was found on a lake surveyed on 
or before 5 July, or if chicks were discovered on the 
lake. Only lakes surveyed after 25 June were used to 
determine productivity. 

To detect changes in the spatial distribution of loons 
over the 12-year period. We divided the study area 
was divided into two regions of differing human pop- 
ulation densities. The Little Susitna River runs east to 
west and divides the study area into southeastern (Was- 
illa, high human population) and northwestern portions 
(Willow, low human population). Lake physiographic 
features are comparable between regions, and lakes 
vary similarly in size, food availability, and shoreline 
topography (Ruggles 1991). 

We used McNemar’s symmetry test for matched 
pairs to compare temporal and geographic differences 
in lake occupancy and productivity (Agresti 1996 pages 
226-229). Alternative hypotheses were constructed to 
indicate a decline in parameter estimates between 1987 
and 1999; no statistical analysis was conducted when 
n <3 lakes. We used P-values to examine strength of 
differences between years and regions. Standardized 
normal standard errors of differences were also cal- 
culated (Agresti 1996 page 228). 


Results 

The number of lakes occupied by any loon or grebe 
species decreased significantly between 1987 and 1999 
across the entire study area (Table 1). Decreases in 
lake occupancy between years were consistent among 
Common Loons, Pacific Loons, and Red-necked 
Grebes (Table 1). Differences in occupancy of lakes by 
loon or grebe pairs, however, were not uniform among 
regions of the Mat-Su Valley. Decreases in the num- 
ber of lakes occupied by any loon or grebe pair and 
by Common Loons were less drastic in the northwest 
region of the study area (Willow) than in the southeast 
region (Wasilla; Table 1), where development is greater. 
Decreases in lake occupancy by Pacific Loons tended 
to be greater in the northwest section (P = 0.1094), 
but conclusiveness of results was hampered by small 
sample sizes (Table 1). Point estimates of decreases in 
occupancy by Red-necked Grebe pairs were similar 
between regions, but variability was much higher in 
the northwest region (Table 1). 

Contrary to lake occupancy, the percentage of lakes 
that fledged Common Loon chicks remained stable 
between 1987 and 1999. Information on productivity 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


of Red-necked Grebes was not collected in 1987, and 
sample sizes were too small to permit analysis of 
changes in productivity for Pacific Loons. Productivity 
of Common Loons was similar on lakes in both regions 
of the study area. 


Discussion 

Temporal and spatial changes in the population dis- 
tribution of loons and grebes in the Mat-Su Valley from 
1987 to 1999 are likely attributable to the concurrent 
rise and spatial settlement of the human population. 
Significant declines in lake occupancy by Common 
and Pacific loons and Red-necked Grebes indicated 
that fewer birds are establishing breeding territories 
at area lakes. Most of the lakes that are no longer used 
by breeding loons and grebes (65%) were located in 
the southeastern portion of the study area — an area that 
has also experienced the greatest human population 
growth. Common Loons have been extirpated from 
lakes within the nearby Municipality of Anchorage, 
and Pacific Loon populations are decreasing there 
(Fair 1998*). Such changes in lake occupancy may 
be indicative of declining loon and grebe populations 
or may reflect large-scale emigration due to the loss 
of suitable nesting habitat. Common Loons inhabiting 
boreal regions of Alaska, however, appear to be stable 
(Groves et al. 1996). 

Population declines of Common Loons and Red- 
necked Grebes throughout North America have oc- 
curred most frequently at the southern boundaries of 
their breeding ranges where habitat quality has become 
marginal owing to development (De Smet 1987; Mc- 
Intyre 1988). In the Mat-Su Valley, a spatial shift in 
lake occupancy from southeast to northwest may be 
linked to habitat quality decreases in the same spatial 
direction. Habitat in the southeastern portion of the 
study area is likely poorer in quality compared to the 
northwest, where fewer people live and more lakes 
remain undeveloped. The stability of lake occupancy to 
the northwest implies that habitat quality is optimal, 
a fact supported by the rapid reoccupation of vacant 
territories (sensu Newton 1998). 

The negative trends in lake occupancy may also 
reflect changes in the demographics of the Mat-Su 
Valley loon and grebe populations. Loons and grebes 
are long-lived species known to return to breed on the 
same lake territories year after year (McIntyre 1988; 
Stout and Neuchterlein 1999). The combination of a 
long lifespan and strong site-fidelity may have caused 
individuals to occupy lakes that had deteriorated in 
quality over several years. When existing occupants 
died or left, the vacancies were not filled, and lake 
occupancy declined over time. Thus, lakes may have 
lost loons and grebes not suddenly through abandon- 
ment by resident breeders, but by failure of new birds 
to replace them after the death of residents. 

Although habitat in the southeastern region may be 
less suitable during selection of a territory, loons may 
remain productive if they choose to nest there. Greater 


2004 MILLS AND ANDRES: CHANGES IN LOON AND GREBE POPULATIONS IN ALASKA ZAS 
TABLE |. Occupancy and productivity of breeding pairs of loons and grebes on lakes in high (Wasilla) and low (Willow) 
human population regions of the lower Matanuska-Susitna Valley, Alaska, in 1987 and 1999. 
Species — stage Percentages of lakes occupied Difference (SE) 
area both years 1987 only 1999 only 1999 — 1987 P-value’ 
Any loon or grebe - occupancy 
entire area (n = 86) 67 Di, 6 -21 (6) 0.0005 
Wasilla (n = 52) 71 29 0 -29 (6) <0.0001 
Willow (n = 34) 62 23 15 -8 (10) 0.2905 
Common Loon - occupancy 
entire area (n = 63) 59 33 8 =25) | + (7) 0.0012 
Wasilla (n = 36) 58 42 0 -42 (8) <0.0001 
Willow (n = 27) 59 22 19 -4 (12) 0.5000 
Pacific Loon - occupancy 
entire area (n = 17) 41 47 12 -35 (17) 0.0547 
Wasilla (n = 10) 40 50 10 -40 (21) 0.1094 
Willow (n = 7) 43 43 14 -29 (26) 0.3125 
Red-necked Grebe - occupancy 
entire area (n = 33) 76 24 0 -24 (7) 0.0039 
Wasilla (n = 25) 76 24 0 -24 (9) 0.0156 
Willow (n = 8) 75 25 0 -25. (15) 0.2500 
Common Loon - productivity 
entire area (n = 34) 24 38 38 OP Fas) 0.5775 
Wasilla (n = 21) 29 33 38 5 (18) 0.5000 
Willow (n = 13) 15 46 38 -8 (25) 0.5000 
Pacific Loon - productivity 
entire area (n = 10) 80 0 20 ~ - 
Wasilla (n = 7) 86 0 14 - - 
Willow (n = 3) 67 0 33 - - 


' one-sided test, 1987 > 1999. 
* small sample sizes precluded reasonable statistical testing. 


breeding success for Common Loons in the south- 
eastern region may reflect differences in the age struc- 
ture of the population. Across numerous species, non- 
breeders and non-territory holders consist mainly of 
sub-adult birds (Smith 1976; Birkhead et al. 1986). 
Because replacements for vacant territories in the north- 
west are likely drawn from the non-territorial cohorts 
(Ruggles 1991), these loons would be expected to have 
lower productivity than experienced territorial loons 
on southeast region lakes. 

Tolerance of human activity by experienced territory 
holders may also explain the differences in Common 
Loon breeding success between the regions. Common 
Loons are known to be relatively flexible in behavior 
and may acclimate themselves to low-level human dis- 
turbance (Titus and VanDruff 1981; Heimberger et al. 
1983; McIntyre 1988). Years of breeding experience 
and gradual habituation to the subtle progression of 
human disturbance would likely result in higher breed- 
ing success for loons that established territories years 
ago. 

Loons may also benefit by nesting in areas where 
humans are moderately active. Recreational activities 
and residents who watch over loons may deter preda- 
tors from taking young or eggs. Reports from Alaska 
Loon Watch volunteers often include stories of resi- 


dents protecting adults and chicks by discouraging 
Bald Eagles and other predators (Tankersley 1987*, 
Mills, personal observation). This interference, com- 
bined with the defensive behavior of an adult loon, 
may be intense enough to dissuade additional preda- 
tion attempts. 

Changes in the distribution of loons and grebes 
warrant concern about the future of these populations 
within the region. The human population is expected 
to continue to increase in the Mat-Su Valley (Alaska 
Department of Natural Resources 1998), which could 
result in further changes to the lake habitat used by 
loons and grebes. Although degradation of lake habi- 
tats are likely widespread across south-central Alaska, 
only a small portion of North America’s populations of 
Common Loon, Pacific Loons, or Red-necked Grebes 
occur in south-central Alaska. People residing in south- 
central Alaska, however, value living with wildlife and 
believe it makes their community “interesting and 
special” (Alaska Department of Fish and Game 2000). 
Therefore, proactive management of lake use and lake- 
shore development, coupled with continued monitoring 
of loon and grebe populations, is needed to improve 
the stability of the freshwater habitats and ensure the 
continued persistence of loons and grebes in the Mat- 
Su Valley of Alaska. 


214 


Acknowledgments 

We sincerely thank Nancy Tankersley and all the 
Alaska Loon Watch volunteers for their devoted inter- 
est and effort. This project was funded by the U. S. 
Fish and Wildlife Service, Region 7 Nongame Migra- 
tory Bird Program, the North American Loon Fund, 
and the Alaska State Parks. Special thanks to Rick 
Mills for assisting in field work. Helpful manuscript 
comments were provided by A. J. Erskine. 


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Received 30 December 2002 
Accepted 2 November 2004 


The Barred Owl, Strix varia in Alberta: Distribution and Status 


LISA TAKATS PRIESTLEY 


Beaverhill Bird Observatory, Box 1418, Edmonton, Alberta T6B 2X3; e-mail: lisa@beaverhillbirds.com 


Priestley, Lisa Takats. 2004 The Barred Owl, Strix varia, in Alberta: Distribution and Status. Canadian Field-Naturalist 118(2): 


215-224. 


Barred Owl distribution and status in Alberta were investigated using over 300 individual records (1912 through 1999) that 
were collected from literature, museum/zoo specimens, nest cards, bird surveys, volunteer raptor banders, and naturalists. Barred 
Owls were distributed throughout much of the boreal forest, aspen parkland, foothill, and mountain ecoregions of Alberta. 
Fifty-four breeding records (46 nests) were found. Barred Owls were associated with older forests and had a clumped distri- 
bution, predominantly along water where larger Balsam Poplar trees provide nesting sites. According to Alberta’s “Status of 
Alberta Wildlife 2000” criteria, the Barred Owl should be assessed as Sensitive. There is no evidence that Barred Owls have 
expanded their range in Alberta in the last 100 years; rather, they have maintained their distribution. 


Key Words: Barred Owl, Strix varia, distribution, status, Alberta, volunteer surveys, nests. 


The Barred Owl, Strix varia, is widely distributed 
throughout North America, ranging from the east 
coast to the western Canadian provinces (American 
Ornithologists’ Union 1998; Johnsgard 1988; Mazur 
and James 2000). It is found from the southern tip of 
Florida to southeastern Alaska. In Canada, the Barred 
Owl is a permanent resident in woodlands of British 
Columbia, through central and western Alberta, central 
Saskatchewan, south-central and southeastern Mani- 
toba, central and southern Ontario, southern Quebec, 
New Brunswick, Prince Edward Island, and the Mari- 
time provinces (Godfrey 1986). There is little pub- 
lished historical information available on the Barred 
Owl in western Canada, except for British Columbia 
(Boxall and Stepney 1982). However, recent studies 
have found this owl to be more common in western 
Canada than originally believed (Campbell et al. 1990; 
Mazur and James 2000; Mazur et al. 1997; Takats 
1998). 

The Barred Owl is still considered rare in northern 
regions of Alberta (McGillivray 1996). It was found 
in the boreal forest region north of Edmonton, in the 
foothills/montane forests of western Alberta, and in 
Jasper National Park. The detailed status and distri- 
bution of the Barred Owl, however, is poorly -docu- 
mented in the province (Boxall and Stepney 1982; 
Kirk and Hyslop 1998). There has been considerable 
debate over the theory that the Barred Owl has only 
recently extended its range into the western states, 
Canada, and Alaska, although their relative rarity in 
the west has been assumed from the paucity of histor- 
ical records (Shea 1973; Taylor and Forsman 1976; 
Leder and Walters 1980; Boxall and Stepney 1982; 
Sharp 1989; Houston and McGowan 1999). It has also 
been suggested that their numbers are increasing in the 
boreal forest due to their increasing tolerance of pre- 
dominantly coniferous forests (Boxall 1986). 

In many parts of its range, the Barred Owl is depend- 
ent on large areas of remote forest with mature and 


old growth trees for nesting, roosting, and foraging 
(Paris 1947; Elody 1983; Devereux and Mosher 1984; 
McGarigal and Fraser 1984; Elody and Sloan 1985; 
Allen 1987; Bosakowski et al. 1987; Johnson 1987; 
Dunbar et al. 1991; James 1993; James et al. 1995; 
Mazur et al. 1997; Mazur et al.1998). In the United 
States it has been reported to nest in interior portions 
of expansive, mature woodland (Allen 1987). The 
typical Barred Owl nest is in a cavity in a large living 
or dead tree or in the top of a broken snag. There are 
records of Barred Owls using more conspicuous plat- 
form nests (i.e., stick nests) built by squirrels or other 
large birds (Mazur et al. 1997), as well as ground nests 
(Robertson 1959; Postupalsky et al. 1997), but these 
are potentially less productive than cavity nests (Pos- 
tupalsky et al. 1997) and are generally quite rare. 

Alberta Environmental Protection (1996) placed 
the Barred Owl on the Yellow B list in the Status of 
Alberta Wildlife report. The Yellow B list includes 
species that are: (1) naturally rare but not in decline, 
(2) naturally rare and have clumped breeding distribu- 
tions, or (3) associated with habitats or habitat elements 
that are, or may be, in decline. The status of Canada’s 
remaining old growth forests is of growing concern 
because of the high rate of harvest (Ellis 1993). In the 
year 2000, the Barred Ow] was placed on the Sensitive 
species list (Alberta Sustainable Resource Develop- 
ment 2000). 

The first step in managing wildlife populations is 
having knowledge of distribution and abundance 
(Mosher and Fuller 1996). The first objective of this 
paper was to compile all historic Barred Owl records 
(published and unpublished) for Alberta, in order to 
determine the past and present distribution in the prov- 
ince. The second, based on recent studies, was to evalu- 
ate this species’ status and general habitat use (as it 
relates to status) in Alberta. The third was to establish 
whether evidence exists to support the suggestion of 
range expansion into the west. 


Pas) 


216 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE 1: Barred Owls banded in Alberta: date, location, age, sex, and name of bander (1966-1999). 


Date Location Age-Sex* 
May 1966 Edmonton L-U 
April 1987 Water Valley SY-U 
November 1988 SSA Ste U-U 
April 1988 Deerland AHY-F 
June 1989 Deerland L-U 
May 1990 Deerland L-U 
May 1991 Niton L-U 
May 1991 Deerland L-U 
May 1992 Deerland L-U 
May 1994 Millarville AHY-F 
June 1994 Tawatina L-U 
June 1994 Uncas L-U 
June 1994 Calling Lake AHY-F 
June 1995 Solomon Creek AHY-F 
August 1996 Gregg Lake AHY-U 
May 1995 Water Valley AHY-F 
May 1995 Water Valley AHY-M 
May 1995 Uncas AHY-F 
May 1995 Uncas L-U 
May 1995 Tawatina L-U 
November 1995 Redwater AHY-M 
November 1995 Se lae2° U-U 
April 1995 Calling Lake AHY-M 
June 1995 Calling Lake AHY-F 
June 1996 Tawatina AHY-F 
June 1996 Tawatina L-U 
May 1996 Calling Lake AHY-M 
May 1996 Calling Lake AHY-M 
May 1996 Calling Lake AHY-F 
June 1996 Calling Lake AHY-M 
July 1996 Calling Lake AHY-M 
August 1996 Calling Lake AHY-M 
May 1997 Tawatina L-U 
May 1998 Calling River L-U 
May 1998 Calling Lake L-U 
June 1998 Uncas L-U 
June 1999 Uncas L-U 
June 1999 Vinca L-U 


* Age — L=nestling/fledgling, AH Y=after hatch year, SY=second year, Sex — U=unknown, M=male, 


Methods 

Information on Barred Owls in Alberta was col- 
lected from a variety of sources, which included: pub- 
lished literature, museum collections (National Museum 
of Canada, National Museum of Natural Sciences (now 
Canadian Museum of Nature), American Museum of 
Natural History, Cornell University Museum of Verte- 
brates, Royal Ontario Museum, University of Calgary 
Museum of Zoology, University of Alberta Museum 
of Zoology, Provincial Museum of Alberta, Alberta 
Breeding Bird Atlas (Federation of Alberta Natural- 
ists), Avian Raptor Nest Cards (Alberta Sustainable 
Resource Development/Beaverhill Bird Observatory 
1999*), banding records (Canadian Wildlife Service 
Bird Banding Office), and personal communications 
with Provincial and National Parks personnel. Records 


Number Bander 


3 E. Jones 
D. Collister 


Pletz/B. Gehlert 
Cromie 

Cromie 
Collister 
Cromie 

Cromie 
Stepnisky/G. Court 
Takats 

Takats 

Collister 
Collister 
Cromie 

Cromie 

Cromie 

Roper 

Halmazna 

Court 
Sissons/G. Court 
Cromie 

Cromie 
Olsen/R. Sissons 
Olsen/R. Sissons 
Olsen/R. Sissons 
Olsen/R. Sissons 
Olsen/R. Sissons 
Olsen/R. Sissons 
Cromie 

Cromie 

Cromie 

Cromie 

Cromie 

Pletz 


BWW PD FB DR OD EOD DBD i OD OD OD OD i i 
Fae Ragen re Ce ee ee eG eee OSs GS eee ee eae 


F=female 


from the following volunteer programs were also col- 
lected: Alberta Bird Records 1983-1988 (Alberta Or- 
nithological Records Committee), Breeding Bird Sur- 
veys, Christmas Bird Counts and May Species Counts, 
volunteer owl survey programs (Edmonton Ow! Prowl, 
Alberta Owl Prowl, Alberta Nocturnal Owl Survey), 
dead raptors turned in to Alberta Sustainable Resource 
Development, and unpublished data including field 
notes. 

All georeferenced Barred Owl sightings and nest 
locations were entered into the Biodiversity Species 
Observation Database (BSOD) (Alberta Sustainable 
Resource Development/Alberta Conservation Associ- 
ation 1998*). Maps comparing distributions from 1912 
to 1975 and 1976 to 1999 were created in ArcView 
(ESRI GIS Mapping and Software). 


2004 


PRIESTLEY: BARRED OWL IN ALBERTA 


Ficure 1. Photo of a Barred Owl pausing before entering a typical nesting cavity (photo by Gordon Court). 


Results 

Of 320 reports of Barred Owls, 297 of these were 
entered into the BSOD database (23 records lacked 
information on date, location, and/or observer). The 
first record of a Barred Owl in Alberta was from a 
specimen collected in 1912, from the Calgary area, 
though this has been considered a mislabeled speci- 
men (Houston and McGowan 1999). The first live 
Barred Owl recorded in Alberta was one heard calling 
along the Athabasca River near Fort McMurray in 
1934 (Preble 1941). 


Dead Owls 

Museum specimens, private collections, and dead 
owls turned in to Fish and Wildlife constituted 53 
Barred Owl locations distributed throughout the prov- 
ince. The most southeasterly report was a dead owl 
turned in to Fish and Wildlife in February 1984 from 
Coronation (Boxall 1986). Morphological information 
collected on 26 owls shows that the majority were 
adults (24 of 26), 12 females, 10 males, 4 unknown 
sex. Boxall and Stepney (1982) reported an unusually 
high number of dead Barred Owls turned in to Fish 
and Wildlife during a short period of time (15 indi- 
viduals from 1982 to 1985). 


Banding, Literature and Personal Communications 
Barred Owls have been banded on 38 occasions in 
Alberta from 1966 to 1999 (CWS Banding Office) by 
thirteen banders (Table 1). There were 15 clutches of 
owls banded at nest sites, as well as nine adult females, 
eight adult males, and three adults of unknown sex. 


Only one Barred Owl was banded before 1987 (2.6 
percent). : 

There were 42 reports of Barred Owls collected from 
publications (Preble 1941; Jones 1956; Jones 1966; 
Salt and Wilk 1958; Salt and Salt 1976; Francis and 
Lumbis 1959; Jones 1987; Sadler and Myers 1976; 
Boxall and Stepney 1982; Rintout and Myers 1983; 
Pintel et al. 1991; Takats 1995) and 63 records collect- 
ed from personal communications. Only 35 of these 
records were pre-1975 (33 percent). Jones (1987) sum- 
marized observations of Barred Owls as being “fairly 
evenly distributed in Alberta, particularly in Alberta’s 
northern forests and heavily forested foothill regions 
of Alberta”. 

Holroyd and VanTighem (1983) report the Barred 
Owl as a rare, year-round resident in Banff and Jasper 
National Parks. They state: 

“Tt occurs regularly near Jasper townsite especially along 

the lower Miette River, where it has nested, and near Cot- 

tonwood Slough (Roy Richards, pers. comm.) and Pyramid 

Lake (R. Salt, pers. comm.). It has been recorded twice 

near North Saskatchewan River Crossing (M. Dyer, pers. 

comm.), twice near Vermilion Lakes Banff, once at Lake 

Annette (K. VanTighem, pers. comm.) and once near Ma- 

ligne Canyon (J. Salt, pers. comm.).” 


Volunteer Surveys 

Breeding Bird Atlas volunteers reported 65 Barred 
Owl locations during a five-year survey from 1987 
through 1991 (data were collected between February 
and July of each year). Atlas data showed that Barred 
Owls were concentrated in the Boreal Forest region, 


218 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE 2. Barred Owl locations recorded on volunteer owl survey programs (EOP — Edmonton Owl Prowl, AOP — Alberta 


Owl Prowl, ANOS — Alberta Nocturnal Owl Survey). 


Date Location Observation Observer 

24 February 1988 Big Lake 1 individual EOP-—_ Beck and Beck (1988) 

5 March 1988 Cooking Lake 1 individual EOP— _ Beck and Beck (1988) 

6 March 1988 Wabamun Creek 1 individual EOP— _ Beck and Beck (1988) 

24 February 1989 Big Lake 1 individual AOP-—_ E. Bamford 

05 March 1989 Cooking Lake 1 individual AOP—_ R. and M. Berg 

06 March 1989 Wabamun Lake 1 individual AOP -—_ S. Jungkind, Belmonte, Gomez, Diener 
21 May 1989 Water Valley 1 individual AOP— _ D. Hutchinson, D. Collister 

10 April 1998 Lac La Biche pair duetting © ANOS — J. Gammon and P. Okrainec 

15 May 1998 Shaw Lake 1 individual ANOS — J. Gammon 

18 April 1998 Hillcrest Road 2 individuals ANOS — M. Heckbert and J. Doll 

26 April 1998 Sibbald 1 individual ANOS — D. Woodsworth, Kanagawa, Mitchell 
27 March 1999 Nojack South 1 individual ANOS — R. Gutsell, R. Wiacek, B. McCulloch 


and primarily coniferous foothills and montane forests 
west of Calgary and in Jasper National Park (Semen- 
chuk 1992). Few records indicated breeding evidence 
and only one possible breeding record was discovered, 
north of Lesser Slave Lake at La Crete along the Peace 
River. During Alberta Christmas Bird Counts from 
1965 to 1998, volunteers in 44 count circles recorded 
53 Barred Owls. Only 17 (32 percent) of these owls 
were recorded before 1990. Only four and two owls 
were recorded during Breeding Bird Surveys and May 
Species Counts, respectively. 

There have been three volunteer surveys conducted 
specifically for owls (Table 2): Edmonton Owl Prowl 
(1988), Alberta Owl Prowl (1989), and the volunteer 
Alberta Nocturnal Owl Survey (1998 and ongoing). 
Seven Barred Owls were found during the first two 
owl surveys (Beck and Beck 1988, personal commu- 
nication). Volunteer Alberta Nocturnal Owl surveyors 
found 13 individual Barred Owls (11 records) on elev- 
en 10 km transects along roads (16 routes were sur- 
veyed throughout the province). The Alberta Bird 
Record contributed 35 more locations of Barred Owls 
in 1988 (Alberta Ornithological Records Committee). 

Two intensive studies have been conducted on the 
Barred Owl in Alberta (Takats 1998; Olsen 1999). 
Takats (1998) conducted a study on distribution and 
abundance of the Barred Owl in the Foothills Model 
Forest (located in the foothills and mountains around 
Hinton and Jasper). Forty-two different territorial 
Barred Owls (10 females, 17 males, and 15 unknown 
sex) were recorded in 1995 and 1996. Of these, seven 
were paired and the other 28 were single but may have 
had mates that did not respond to broadcasts (Takats 
1998). Surveys were continued in 1997 and 1998 and 
four additional Barred Owls were located. The density 
of Barred Owls was determined to be 0.05 and 0.04 
owls/km?, in 1995 and 1996, respectively. 

The habitat used by Barred Owls at 45 calling loca- 
tions in the Foothills Model Forest was predominant- 


ly older mixedwood forest containing Trembling Aspen 
(Populus temuloides), White Spruce (Picea glauca) 
and Balsam Poplar (Populus balsamifera). Roosting 
and foraging occurred in a variety of stand types, but 
were also predominantly older mixedwood Populus 
sp. and spruce (Takats 1998). 

A study to evaluate the effects of forest fragmenta- 
tion on bird communities was initiated in north-central 
Alberta (Schmiegelow and Hannon 1993). Preliminary 
work located six Barred Owl territories, and evidence 
of three breeding pairs was discovered in the Calling 
Lake area (G. Court, personal communication). Two 
additional territories and four additional breeding 
sites were found in a study from 1996 through 1998 
(Olsen 1999). During the breeding season, the density 
of Barred Owls was 0.04 pairs/km?. Owls were found 
to use old growth mixed forest stands (Olsen 1996). 


Breeding Records 

There were 54 breeding records (only three found 
before 1975). The first evidence of breeding was re- 
corded in 1949 (Grant 1966), but the first nest was 
not discovered until 1966, in Edmonton (Jones 1966) 
(Table 3). Boxall and Stepney (1982) reported eight 
breeding records, and Semenchuk (1992) reported six 
confirmed breeding records during the Provincial 
Breeding Bird Atlas Project (1987-1991). The north- 
ern-most breeding record was a nest found at 58° 19' 
latitude and 116° 17' longitude near La Crete (Takats 
1995). Some local residents have reported Barred Owls 
to be present for over 20 years. 

Breeding records for Barred Owls were not com- 
mon (n = 54) and only 46 nests have been found (1966 
through 1999): 38 (82.6 percent) were in natural cavi- 
ties (or bowls on top of dead trees that had broken off), 
two (4.3 percent) were in stick nests, one (2.2 percent) 
was in a man-made stick nest, and five (10.9 percent) 
were in nest boxes. In most cases owls nested in older 
mixedwood stands and used large diameter Populus 
sp. trees (> 40 cm) (primarily Balsam Poplar) for 


2004 


TABLE 3: Confirmed breeding records of Barred Owls for Alberta. 


Man-Made Platform 


Date Location Nest Type 
1949 Lesser Slave Lake Unknown 
1966 Edmonton Cavity Nest 
1968* Edmonton Cavity Nest 
1976* Blue Lake Cavity Nest 
LOTA* Jasper Cavity Nest 
1977* Blue Lake Cavity Nest 
1978 Chilver Lake Unknown 
IO Miette River Cavity Nest 
1980* Miette River Cavity Nest 
1984** Spruce Grove Cavity Nest 
1988 Deerland Cavity Nest 
1989 Deerland Cavity Nest 
1990 Deerland Cavity Nest 
199] Deerland Cavity Nest 
199] Niton Cavity Nest 
1992 Deerland Cavity Nest 
1993 La Crete Cavity Nest 
1994 La Crete Cavity Nest 
1994 Millarville Unknown 
1994 Uncas 

1994 Tawatinaw Cavity Nest 
1994 Elk Island Stick Nest 
1994 Calling Lake Unknown 
1994 Miette River Cavity Nest 
1995 Lynx Creek Cavity Nest 
1995 Miette River Cavity Nest 
1995 La Crete Cavity Nest 
1995 Tawatinaw Cavity Nest 
1995 Cross Lake Cavity Nest 
1995 Uncas Nest Box 
1995 Calling Lake Cavity Nest 
1995 Calling Lake 2 Cavity Nest 
1996 Bragg Creek Cavity Nest 
1996 Solomon Creek Cavity Nest 
1996 Blackcat Ranch Cavity Nest 
1996 Miette River Cavity Nest 
1996 Tawatinaw Nest Box 
1996 Calling Lake Unknown 
1997 Cross Lake Cavity Nest 
1997 Miette River Cavity Nest 
1997 Tawatinaw Cavity Nest 
1997 Lac La Biche Stick Nest 
1997 Calling Lake Cavity Nest 
1998 Uncas Cavity Nest 
1998 Solomon Creek Cavity Nest 
1998 Calling River Cavity Nest 
1998 Calling Lake Cavity Nest 
1999 Uncas Nest Box 
1999 Tawatinaw Nest Box 
999 Fe Vinca Cavity Nest 
1999 Cross Lake Nest Box 


PRIESTLEY: BARRED OWL IN ALBERTA 219 


Observer/Reference 


Grant (1966) 

Jones (1966) 

E. Jones (personal communication) 
Boxall and Stepney (1982) 

Boxall and Stepney (1982) 

Boxall and Stepney (1982) 

Wiseley (personal communication) 
Richards (personal communication) 
Richards (personal communication) 
R. Copeland (personal communication) 
R. Cromie (Nest Card) 

R. Cromie (Nest Card) 

R. Cromie (Nest Card) 

R. Cromie (personal communication) 
H. Pletz/B. Gehlert 

R. Cromie (Nest Card) 

Takats (1995) 

Takats (1995) 

D. Collister (personal communication) 
R. Cromie (Nest Card) 

R. Cromie (Nest Card) 

H. Pletz (personal communication) 
G. Court (personal communication) 
Takats (1998) 

Takats (1998) 

Takats (1998) 

A. Miller (personal communication) 
R. Cromie (Nest Card) 

R. Cromie (Nest Card) 

R. Cromie (Nest Card) 

G. Court (personal communication) 
G. Court (personal communication) 
H. Pletz (personal communication) 
Takats (1998) 

Takats (1998) 

Takats (1998) 

R. Cromie (Nest Card) 

Olsen (1999) 

A. Karvonen (personal communication) 
L. Takats (personal communication) 
R. Cromie (Nest Card) 

Olsen (1999) 

Olsen (1999) 

R. Cromie (personal communication) 
L. Takats (personal communication) 
T. Roper (Nest Card) 

Olsen/Cromie (Nest Card) 

R. Cromie (Nest Card) 

R. Cromie (Nest Card) 

H. Pletz (Nest Card) 

R. Cromie (Nest Card) 


* Boxall and Stepney (1982) ** Nest cut down, two eggs cracked, one survived, owl held at Valley Zoo, Edmonton 


*** One young Barred Owl was fostered in to this nest. 


nesting (Takats 1998; Avian Raptor Nest Cards; Olsen 
1999). Nest boxes built specifically for Barred Owls 
were readily taken over in areas where natural cavities 
were not abundant (G. Court, personal communica- 
tion; R. Cromie, personal communication). 


Distribution 

Most occurrences of Barred Owls were in the boreal 
forest, foothill, and mountain ecoregions from 1912 
through 1974 (Figure 2) and from 1975 to the present 
(Figure 3). Few owls were recorded in the parkland, 


220 THE CANADIAN FIELD-NATURALIST Vol. 118 


Barred Owl Occurrences 
1912-1974 


Medicine Hat 


FIGURE 2. Barred Owl distribution from 1912 through 1974 (map courtesy of Alberta Conservation Association/Alberta Sus- 
tainable Resource Development). 


NR 
i) 


2004. PRIESTLEY: BARRED OWL IN ALBERTA 


Barred Owl Occurrences 
1975-1999 


FiGuRE 3. Barred Owl distribution from 1975 through 1999 (map courtesy of Alberta Conservation Association/ Alberta 
Sustainable Resource Development). 


222 


and only one individual was recorded in the northern 
limit of the grassland ecoregion. Detection of Barred 
Owls appears to have increased after 1975, but there 
have been only two records outside the range of the 
early distribution map. 


Discussion 

The Barred Owl is distributed over much of Alberta’s 
forested area. Evidence suggests that Barred Owl dis- 
tribution in the province has changed little over the 
last 100 years. As more detailed studies are conducted, 
as there is an increase in the number of interested 
naturalists exploring isolated woodlands, and as there 
is increased access to remote areas, more reports of 
Barred Owls will accumulate, Oeming (1957) and 
Jones (1966) concur. Mazur et al. (2000) suggested that 
Barred Owls are shy of humans, which may account 
for their secretive nature. 

In British Columbia the first records of Barred Owls 
were from Liard Crossing in 1943 and Nechako Low- 
lands in 1946 (Campbell et al. 1990). Scotter et al. 
(1985) reported a Barred Owl record from 1977 in 
the South Nahanni River, Northwest Territories, which 
shows that Barred Owls have existed northwest of 
Alberta for over 20 years. As well, the idea that the 
Barred Owl is adapting to the coniferous boreal forest 
does not hold true. Barred Owls have shown a clumped 
distribution in areas that contain large deciduous trees 
(predominantly riparian areas). They use Balsam Pop- 
lars for nesting and rely on a more open subcanopy 
for flight, although the roost cover provided by White 
Spruce trees is important, particularly in the winter 
(Takats 1998). Some of the first records of Barred 
Owls occurred along riparian areas. 

The Sensitive designation assigned by Alberta Sus- 
tainable Resource Development (2000) is well founded. 
Based on recent studies and personal communications 
with raptor banders and naturalists, the Barred Owl 
should not be considered rare in Alberta; however, 
this species does have a clumped breeding distribu- 
tion. As well, these owls rely on cavities in old, large 
diameter Populus sp. trees for nesting, and select old 
and/or mature mixedwood forests (a habitat that is in 
decline) to fulfill their life requisites (Takats 1998; 
Olsen et al. 1996*; Olsen 1999; Mazur and James 2000). 

As older forests are usually targeted first for har- 
vest, the amount of old growth forest remaining de- 
creases and stands become increasingly fragmented. 
Loss of nesting, roosting, and foraging habitat occurs 
when a forest is clearcut; moreover, Great Horned 
Owls, Bubo virginianus, are known to favour frag- 
mented landscapes. There is direct conflict between 
these two owl species, with the Barred Owl losing out 
to the larger Great Horned Owl (Bent 1961; Bosa- 
kowski 1994; Laidig and Dobkin 1995; Takats 1998 
and field notes; Olsen 1999; G. Court, personal com- 
munication). Barred Owl numbers and range are lim- 
ited by the amount of adequate nesting habitat avail- 
able where they can reproduce successfully and fledge 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


their young without interference from competitors 
and predators. 


Acknowledgments 

Funding for parts of this work was provided by 
Foothills Model Forest, Alberta Sport, Recreation, 
Parks and Wildlife Foundation, Alberta Sustainable 
Resource Development (Fish and Wildlife), Environ- 
ment Canada (Canadian Wildlife Service), Beaverhill 
Bird Observatory, and Weldwood of Canada, Hinton. 
The Bird Banding Office (Canadian Wildlife Service) 
provided Barred Owl banding information for Alberta. 
The Federation of Alberta Naturalists provided loca- 
tions from the Breeding Bird Atlas. The Alberta Con- 
servation Association/Alberta Sustainable Resource 
Development provided use of the Biodiversity Species 
Observation Database (BSOD) to create maps of the 
distributions. 

I thank Tony Erskine, Jim Duncan, and Bob Nero 
for reviewing the submitted manuscript. Gordon Court, 
Jim Beck, and Geoff Holroyd initially reviewed the 
manuscript, provided owl location information, and 
continue to encourage me with my work in raptor 
ecology. Lance Engley (Alberta Conservation Asso- 
ciation) provided the maps. Chuck Priestley, Dawn 
Birn, Kris Kendell, and Jeff Adamyk also provided 
some editorial comments. Jim Duncan and Kurt Mazur 
provided the idea to write the paper in the first place. 
I acknowledge Jim and Barb Beck (the ‘Owl Nuts’) 
for providing the Edmonton and Alberta Owl Prowl 
data as well as many other locations. Edgar Jones, 
Doug Collister, Ray Cromie, Wayne Smith, Hardy 
Pletz, Bob Gehlert, Ben Olsen, Trevor Roper, and 
Rick Lauzon (AXYS) provided many owl locations. 
Thanks also to Wayne Roberts (University of Alberta 
Museum of Zoology), Michel Gosselin (National 
Museum of Canada [now Canadian Museum of Na- 
ture), Mark Steinhilber (Provincial Museum of Alber- 
ta), and Rusty Copeland (Valley Zoo, Edmonton) for 
providing information on museum and zoo specimens. 
Thanks to Howard Troughton for providing the Christ- 
mas Bird Count information in a ready-to-use file. 
Special thanks to Stephen Glendinning and Jeff 
Adamyk, who spent many hours conducting broad- 
cast surveys for owls in the foothills. Finally, thank you 
to all the volunteer raptor banders, owl surveyors, and 
raptor enthusiasts who have contributed most of the 
information we know about Barred Owls in Alberta. 


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Received 30 July 2002 
Accepted 22 June 2004 


Use of Eelgrass, Zostera marina, Wrack by Three Species of Ladybird 
Beetles (Coleoptera: Coccinellidae) in Prince Edward Island 


DaviD J. GARBARY, SARAH FRASER, CARRIE FERGUSON, and RANDOLPH F. LAUFF 
Department of Biology, St. Francis Xavier University, Antigonish, Nova Scotia B2G 2WS5 Canada 


Garbary, David J., Sarah Fraser, Carrie Ferguson, Randolph F. Lauff. 2004. Use of Eelgrass, Zostera marina, wrack by three 
species of ladybird beetles (Coleoptera: Coccinellidae) in Prince Edward Island. Canadian Field-Naturalist 118(2): 
225-228. 


Large numbers of the introduced ladybird beetle, Coccinella septempunctata L., were present at Wood Islands and Green 
Point, Prince Edward Island, in wrack consisting primarily of Zostera marina L. (Eelgrass). The wrack occurred in a 0.5 to 
1.0 m band parallel to the shore, and was from five to 25 cm thick. The other ladybirds, Propylea quatuordecimpunctata 
(L.), an introduced coccinellid also found in high numbers, and an individual of the native Hippodamia tredecimpunctata 
(Say) were found only at Wood Islands. At both sites the ladybird beetles occurred in the mid-intertidal zone along at least 
100 m of shoreline, and were absent to rare on the terrestrial vegetation above the high tide mark. At four of the other eight 
sites surveyed, occasional individuals were present in the wrack, but they were no more abundant than could be observed on 
landward vegetation. Mean densities of C. septempunctata at the two primary sites were 52 m? (Green Point) and 410 m7? 
(Wood Islands). 


Key Words: Coccinella septempunctata, Propylea quatuordecimpunctata, Hippodamia tredecimpunctata, ladybird beetles, 


Coccinellidae, Zostera marina, Eelgrass, intertidal zone, Prince Edward Island. 


With over 150 species and subspecies, the Coc- 
cinellidae of Canada and Alaska are a conspicuous 
and ecologically important element of the terrestrial 
biota (McNamara 1991). In addition to the native 
fauna, there is great interest in the distribution of 
invasive ladybirds and the subsequent loss of native 
biodiversity (Gordon and Vandenberg 1991). This is 
also true in eastern Canada where Hoebeke and Wheel- 
er (1996) and McCorquodale (1998) and Majka and 
McCorquodale (in press) reported the spread of intro- 
duced species in the Maritimes. 

Despite being a terrestrial family, there are several 
reports of Coccinellidae from beaches and salt marshes 
in both freshwater and marine environments (Davis 
and Gray 1966; Schaefer et al. 1987; Turnock 1996; 
Pupedis 1997; review by Nalepa et al. 1998). Here we 
report on the mass occurrence of living ladybird beetles 
in the marine intertidal of Prince Edward Island and 
their association with Eelgrass (Zostera marina) wrack. 


Materials and Methods 

Ten intertidal locations on Prince Edward Island 
(Table 1) were visited on 30-31 August 2002 for the 
examination of Eelgrass wrack as part of a study on 
Eelgrass decline. An abundant population of ladybird 
beetles was at site one (Wood Islands), and several hours 
were spent photographing and quantifying the assem- 
blage. At subsequent sites three individuals inspected 
the wrack for a minimum of 15 min along at least 100 m 
of shoreline. If only a few beetles were observed, their 
presence was noted. If many were present, densities 
were calculated using a 20 x 20 cm quadrat. The quad- 


rat was placed haphazardly on the eelgrass wrack (n = 
25) at irregular intervals on the shore. 

Characteristics of the primary study sites were: The 
Wood Islands site was a sandy beach with sandstone 
outcrops. The primary species of seaweeds associated 
with the wrack were recorded. Green Point was a salt 
marsh adjacent to an abandoned wharf, and common 
plants in the marsh were recorded. At Green Point the 
wrack was almost exclusively Eelgrass. Characteris- 
tics of other sites are mentioned, where necessary, in 
the body of the text. The weather on both days was 
mostly bright and sunny; however, a mild rain fell at 
Marchwater (1800 h, 30 August) and Belmont Prov- 
incial Park was extremely windy (0800 h, 31 August). 
Coordinates of primary sites were determined using a 
global positioning system (Garmin GPS 12, Olanthe, 
Kansas; Table 1). 

Beetles were identified using the keys in Gordon 
(1985) and by comparison with specimens in the in- 
sect collection at St. Francis Xavier University. Voucher 
specimens have been deposited in the Herbarium of 
St. Francis Xavier University (STFX) and in the insect 
collection. 


Results 

Three species of ladybird beetles, Coccinella sep- 
tempunctata, Hippodamia tredecimpunctata, and Prop- 
ylea quatuordecimpunctata, were present in intertidal 
Eelgrass wrack (Zostera marina) in Prince Edward 
Island during late August. All species were present at 
Wood Island; however, only C. septempunctata was 
collected from the other sites. 


PIT) 


226 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE 1. Relative abundance of P. quatuordecimpunctata, H. tredecimpunctata and C. septempunctata in sites across Prince 
Edward Island, and their corresponding GPS coordinates. “Low” corresponds with an observation of < 5 individuals. 


Site Coordinates 
Wood Islands 45°56'N 
62°45'W 
St. Peters Bay 46°26'N 
62°28'W 
March Water 46°29'N 
63°44'W 
Belmont Provincial Park 46°31'N 
63°49'W 
Green Point 46°35'N 
63°52'W 
Cascumpec Bay 46°45'N 
64°04'W 
Linkletter Provincial Park 46°24'N 
63°51'W 
Victoria Provincial Park 46°12'N 
63°30'W 
West River Bridge 46°11'N 
63°14'W 
Pinnette Provincial Park 46°04'N 
62°54'W 


At Wood Islands, C. septempunctata occurred pri- 
marily in a loose to dense band of Eelgrass wrack, 
0.2-1.0 m wide, that formed in the mid-intertidal zone 
on a sandy beach (Figure 1). No coccinellid beetles 
were found in the dry wrack in the splash zone nor 
in the mixed herbaceous vegetation above the high 
tide mark. A few beetles were also found in small 
clumps of wrack lower in the intertidal zone that had 
recently been inundated by the incoming tide and 
very gentle wave action. The wrack consisted mostly 
of leaves of Eelgrass and was mixed with several com- 
mon seaweeds including the brown algae, Fucus ser- 
ratus, Chorda filum, Chordaria flagelliformis, and the 
red algae, Chondrus crispus (Irish moss), Palmaria 
palmata (Dulse) and Furcellaria lumbricalis. The 
Eelgrass was relatively fresh and had numerous green 
leaves. 

The beetles were present on the exposed wrack to 
about two cm into the wrack mass in loosely packed 
leaves (Figure 2). Beetle density was 410 + 340 m? 
(mean + SD). Beetles were absent deep within the 
20-30 cm thick wrack bundles. The beetles walked 
along the surface of the leaves and were single or in 
clumps of two-five individuals. Occasional beetles 
were present on the bare sand adjacent to the wrack 
or on clumps of seaweed (mostly Chorda filum and 
Fucus serratus) that separated from the main body of 
wrack. No flying beetles were observed; however, 
walking beetles occasionally spread their elytra. 

At Green Point, the Eelgrass wrack accumulated 
in a 0.5 —2 m band in the mid-intertidal zone of a salt 
marsh. This was a typical marsh dominated by Spar- 
tina alterniflora in the lower marsh and a mixture of 


Abundance 


Very high 
Absent 
Absent 
Absent 

High 
Absent 
Low 
Low 
Low 


Low 


S. patens, Scirpus americanus, S. maritimus, Glaux 
maritimus, Triglochin maritima and Limonium nashii 
in the mid-intertidal zone. At various sites along the 
shore there was extensive accumulation of old Eelgrass 
in the upper intertidal zone. The Eelgrass wrack tend- 
ed to form a blanket that matted down the surround- 
ing vegetation or occasionally was suspended up to 
30 cm above the ground. 

The beetles (all C. septempunctata) were found on 
the Eelgrass in the mid-intertidal zone, with rare indi- 
viduals on the surrounding grasses and sedges. The 
insects were common (52 + 8.5 m7) over more than 
100 m of shoreline, and were typically single. Beetles 
were absent on the upper intertidal wrack, and none 
were observed in the lower intertidal zone. The insects 
walked on the Eelgrass and did not fly. In addition to 
the ladybird beetles, one Leptinotarsa decemlineata 
(Colorado Potato Beetle) was observed on the wrack. 

At Linkletter and Pinnette Provincial Parks and West 
River Bridge, 3-5 beetles (all C. septempunctata) 
were found on Eelgrass in the upper intertidal zone. 
These sites were primarily sand beaches with either 
scattered clumps or continuous carpets of Eelgrass up 
to 2 m wide in the high intertidal zone. At St. Peters 
Bay, Belmont and Victoria Provincial Parks and Cas- 
umpec Bay, beetles were absent, despite the presence 
of extensive Eelgrass wrack. 

Upon our return to Wood Islands, 28 h after the 
initial observations, we found that the previous day’s 
wrack had largely disappeared. A few hundred dead 
ladybird beetles were present in the remaining wrack 
and on sandstone outcrops along with a few live 
individuals. 


2004 


GARBARY, FRASER, FERGUSON, and LAUFF: USE OF EELGRASS BY LADYBIRD BEETLES 


Ficure |. Ladybird beetle site at Wood Islands, Prince Edward Island, showing band of intertidal wrack on beach with investi- 
gator (C.F.) examining Eelgrass. 


Discussion 

Although the records for both introduced species 
(C. septempunctata, P. quatuordecimpunctata) do not 
represent range extensions, it is disturbing that they 
did make up essentially all of the coccinellid fauna at 
the sampled sites. P. quatuordecimpunctata has been 
recorded in PEI since at least 1994 (Hoebeke and 
Wheeler 1996). The first records for C. septempunc- 
tata on PEI (Charlottetown) are from 1982; it is now 
abundant in the province (Majka and McCorquodale; 
in press). The two sites where C. septempunctata was 
abundant shared the feature of having abundant Eel- 
grass wrack in the mid-intertidal zone. The beetles 
were generally not found in the older, dry wrack at the 
upper part of the shore nor in the vegetation above 
the high tide mark (Wood Islands and Green Point), 
and the surrounding salt marsh vegetation (Green 
Point). Examination of the Eelgrass substratum showed 
no conspicuous populations of invertebrates that might 
be a suitable food source. In addition, the beetles were 
sluggish, suggesting exhaustion due to struggling or 
partial suffocation. The location of the beetles and their 
behaviour suggest that the they were only recently 
washed up to the eelgrass wrack, a substrate which 
allowed them firm footing. Sites with few or no beetles 
had less fresh Eelgrass wrack. The intertidal substra- 
tum at these sites tended to be rockier, with better 
developed populations of seaweed. 

Although terrestrial, both C. septempunctata and 
Propylea quatuordecimpunctata have previously been 
found in coastal habitats including salt marshes in 


Connecticut, North Carolina and Delaware (Schaefer 
et al. 1987; Pupedis 1997; Nalepa et al. 1998). Davis 
and Gray (1966) also reported another ladybird beetle, 
Naemia serriata, from a salt marsh in North Caro- 
lina. The mass occurrences of C. septempunctata that 
we found are different from that in Delaware in which 
numerous dead and some living individuals were 
washed ashore following deposition in the ocean and 
association with seaweed wrack (Schaefer et al. 


a 
2. 


Coccinella septempunctata on mid-intertidal 
Eelgrass wrack. Note: The white spots on the leaves are a 
calcified red alga. 


FIGURE 


228 


1987). The North Carolina mass occurrence is similar 
to that reported by Schaefer et al., and involved mass 
mortality and shore deposition (Nalepa et al. 1998). 
Turnock (1996) suggested that accumulations of lady 
beetles on the shores of Lake Manitoba resulted from 
wave deposition and subsequent migration up the 
shore to structures protruding from the sand. Given 
the localization of C. septempunctata in this study, 
particularly at Green Point, the Eelgrass seems to have 
been a fortunate substratum on which the beetles could 
take hold, and rest prior to dispersal. 

Although there were conspicuous corpses of lady- 
bugs in the wrack at Wood Islands on the day after the 
survey, the vast majority of the population had dis- 
appeared from the shore. There was no evidence that 
the beetles had moved onto the grassy field adjacent 
to the beach. Invasions of Coccinellidae into North 
America show that these species can cover a large dis- 
tance (Schaefer et al. 1987). McCorquodale (1998) 
gives range expansion rates of 31-440 km y"! across 
North America for four introduced coccinellids in 
Nova Scotia, although he concedes that not all of this 
is likely to be long distance flights. Schaefer et al. 
(1987) comment on the occurrence of C. septempunc- 
tata on Sable Island, 300 km from mainland Nova 
Scotia. Thus the 20 km flight across the Northumber- 
land Strait to or from Nova Scotia is reasonable, espe- 
cially if a suitable wind is available. 

Our records of C. septempunctata on the north and 
south shores of PEI are consistent with eelgrass pro- 
viding a staging substratum during migration. Migra- 
tions of Hippodamia convergens cover hundreds of 
kilometres in single flights in California from San 
Francisco and the Imperial Valley to the Sierra Nevadas 
(Hodek 1973). However, the aggregations we observed 
in Prince Edward Island are more likely reflective of 
accidental downing of migrating populations as a result 
of weather conditions, followed by subsequent wash- 
up. The Eelgrass provided a better refugium than other 
substrata in the mid-intertidal zone. We suggest that 
the beetles are only saved by the Eelgrass wrack, and 
that there is no particular attraction to it as previously 
described for similar vegetation with C. septempunc- 
tata (Schaller and Nentwig 2000; Frantsevich and 
Zolotov 2001). 


Acknowledgments 

We thank Christine Nalepa and Chris Majka who 
provided helpful discussion and literature; the com- 
ments of two anonymous reviewers were also an asset 
to this paper. Barry Taylor kindly assisted with plant 
identification and David McCorquodale verified iden- 
tifications of beetles in our insect collection. This 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


work was supported by a grant from the Natural Sci- 
ences and Engineering Research Council (NSERC) 
to D. J. G., funding from the Nova Scotia Museum to 
R. FE. L., and by NSERC undergraduate awards to S. 
F and C. F. 


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Received 17 September 2002 
Accepted 15 November 2004 


Predicting the Effects of Cerulean Warbler, Dendroica cerulea 
Management on Eastern Ontario Bird Species 


JASON JoNES!, WILLIAM J. MCLEISH, and RALEIGH J. ROBERTSON 


Department of Biology, Queen’s University, Kingston, Ontario K7L 3N6 Canada 
| Present address, Department of Biology, Vassar College, Poughkeepsie, New York 12604 USA; e-mail: jajones @ 
vassar.edu 


Jones, Jason, William J. McLeish, and Raleigh J. Robertson. 2004. Predicting the effects of Cerulean Warbler, 
Dendroica cerulea, management on eastern Ontario bird species. Canadian Field-Naturalist 1 18(2): 229-234. 


Single-species habitat management strategies are often undertaken without explicit consideration of their effects 
on the larger community. Here we explore the potential effects of managing eastern Ontario deciduous forests 
for the Cerulean Warbler (Dendroica cerulea) by examining its potential as a biodiversity indicator species and 
as an umbrella species. Our results indicate that the Cerulean Warbler would not be an effective biodiversity 
indicator, as its distribution across the studied landscape did not coincide with areas of high avian species rich- 
ness. However, the Cerulean Warbler may be effective as an umbrella species for the maintenance of populations 
of other canopy-nesting species that require mature deciduous forest habitats. It is hoped that the conclusions 
reached in Ontario, while perhaps not directly transferable to all parts of the breeding range, encourage other 
Cerulean Warbler researchers to ask similar questions in their study areas. 


Key Words: Cerulean Warbler, Dendroica cerulea, conservation utility, biodiversity indicator species, umbrella 


species, eastern Ontario. 


Few North American songbirds are receiving the 
scientific and conservation attention that is currently 
focused on the Cerulean Warbler (Dendroica cerulea; 
Robbins et al. 1992; Rosenberg et al. 2002*; Hamel et 
al. 2004). The Cerulean Warbler has suffered signifi- 
cant long-term breeding population declines (annual 
declines of 3.04% over 1966-2000; Link and Sauer 
2002) which are largely attributed to habitat destruction 
on both the breeding and wintering grounds (Robbins 
et al. 1992). Concern over the long-term health of this 
species has led to its designation as threatened, rare, or 
of special concern in the United States and as a Species 
of Special Concern in Canada (Robbins et al. 1992; 
Hamel 2000; COSEWIC 2003*). 

In two separate meeting in 2001 and 2002, a group 
of academic and governmental scientists, land mana- 
gers, industry biologists and non-governmental organi- 
zations formed the Cerulean Warbler Technical Group 
(CWTG): an effort to develop a proactive, broad-based, 
multiple stakeholder approach to Cerulean Warbler 
conservation in both North and Latin America (Hamel 
et al. 2004). One of the outcomes of these meetings 
was a breeding grounds research plan that is designed 
to identify key population limitation factors and ex- 
plore forest management options for the maintenance 
and creation of Cerulean Warbler habitat on a range- 
wide basis. 

Ironically, the very thing that limits our ability to 
currently diagnose specific reasons for breeding ground 
population declines — a general lack of natural and life- 
history information — may also limit the efficacy of 


the CWTG research plan. Here, we take advantage of 
data collected during one of the longest running Ceru- 
lean Warbler research programs in North America 
(dating to 1994; Oliarnyk and Robertson 1996; Jones 
et al. 2000, 2001, 2004; Jones and Robertson 2001: 
Barg et al. 2005) to make predictions regarding the 
potential effects of managing deciduous forests for 
Cerulean Warblers on sympatric bird species in eastern 
Ontario. Specifically, we ask two questions. One, will 
the promotion of Cerulean Warblers and Cerulean 
Warbler habitat promote the preservation of avian 
diversity? In other words, is the Cerulean Warbler a 
biodiversity indicator (sensu Landres et al. 1988)? 
Two, can the Cerulean Warbler act as an umbrella 
species for other bird species with similar life — and 
natural histories? The protection of the habitat of the 
umbrella species ideally results in the protection of 
the habitat of those species whose requirements are 
subsumed by those of the umbrella (Launer and Mur- 
phy 1994; Berger 1997; Simberloff 1998). 


Study Area and Methods 
This investigation was conducted at the Queen’s 
University Biological Station (QUBS), Ontario 


(44°34'N, 76°20'W), within the Great Lakes-St. Law- 
rence mixed forest region. Our study area was restricted 
to approximately 2 600 ha of research tracts managed 
by QUBS. The landscape in the area is dominated by 
mature, secondary-growth, lowland mixed deciduous 
forest, interspersed with rocky outcrops, marshes, lakes 
and abandoned agricultural fields. 


229 


230 


Bird surveys. In 1997 and 1998, we surveyed birds 
on QUBS property using variable-circular-plot point 
counts (Reynolds et al. 1980; Jones et al. 2000). We 
surveyed 80 stations in 1997 (17 May — 20 June) and 
67 stations in 1998 (21 May — 21 June); the 1998 sta- 
tions were a subset of those sampled in 1997. The 
point-count stations were located to maximize spatial 
coverage of the study area. Each point count was 
10 min long. Each station was separated by at least 
200 m to minimize the potential for double-counting 
individuals. Point counts were conducted between 
0.5 hr before sunrise and 3 hr after sunrise EST in 
order to sample during peak song activity, and were 
only conducted under calm weather conditions. For our 
analyses we included birds detected within 100 m of 
the plot center. Probability of detection was similar for 
all species analysed, as the detection thresholds for all 
but one species (Black-and-white Warbler, Mniotilta 
varia, 92 m) were beyond 100 m (Jones unpublished 
data). Data from the first two visits per station each 
year were used in the analyses to facilitate comparison 
between years and because two visits are sufficient to 
confirm the presence or absence of Cerulean Warblers 
(Jones et al. 2000). 


Vegetation Surveys. We collected vegetation data 
at 59 point-count stations in 1997 at five circular sub- 
plots, each with a radius of 5 m. The first subplot was 
centered on the point count station center and the other 
four were located 50 m away in each of the cardinal 
directions. Within each of the subplots we counted the 
number of saplings [stems < 3.0 cm diameter at breast 
height (dbh)], measured the dbh of all stems 2 3.0 cm 
and grouped them into two size classes (3.0 — 15.0 cm 
dbh, 15.0 — 30.0 cm dbh). Using an imaginary | m radius 
cylinder projected upward from the forest floor, we 
estimated cover within 3 m height intervals from the 
ground to the top of the canopy. Total cover and per- 
cent cover of each woody plant species was estimated 
by eye in each height interval on a scale of 0 to 10 
(0 = 0% cover, 10 = 100% cover). Two observers made 
all cover estimates; the two observers spent several 
days prior to data collection assuring that their cover 
estimates were within 10% of one another. For analysis 
we reduced these cover estimates to two variables: 
maximum cover below 6 m (understory cover) and 
maximum cover above 12 m (canopy cover). Each 
vegetation variable was averaged across subplots to 
describe the habitat of the point-count station. We 
measured only those vegetation variables thought to 
be important to Cerulean Warblers; the importance of 
these variables has been supported by subsequent re- 
search (Jones and Robertson 2001; Jones et al. 2001). 
We also restricted the number of vegetation variables 
for analytical reasons (see below). 


Data Analysis. In our analyses, we included only 
those species known to breed in our study area. We also 
excluded species that are not adequately sampled by 
diurnal point counts, such as colonial nesters (e.g., 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Barn Swallow, Hirundo rustica), nocturnal species (e.g., 
Whip-poor-will, Caprimulgus vociferus), waterfowl 
(e.g., Wood Duck, Aix sponsa), and waders (e.g., Great 
Blue Heron, Ardea herodias). 


(1) Biodiversity indicator evaluation 

We tested whether the presence of Cerulean War- 
blers was a predictor of overall bird species richness. 
For the purposes of these analyses, we defined spe- 
cies richness as the number of species detected in the 
first two visits to a point-count station, excluding the 
Cerulean Warbler if present. We used randomization 
tests to compare the mean species richness at stations 
where Cerulean Warblers were present to the expect- 
ed species richness at a randomly generated sample 
of points (Chase et al. 2000). In these tests, the mean 
species richness was calculated for a random sample 
of point-count stations, with the number of random 
stations equaling the number of stations where Ceru- 
lean Warblers were detected in a given year. We iter- 
ated this procedure 1000 times to generate an expected 
distribution of mean species-richness values. We then 
compared the observed species richness associated 
with Cerulean Warblers and determined its statistical 
significance. These randomization tests were performed 
using S-PLUS 4.0 (Mathsoft 1997*). Values reported 
in the results are means + SE. 


(2) Umbrella species evaluation 

The initial step in evaluating the potential of the 
Cerulean Warbler as an umbrella species was to estab- 
lish an ecological context. We categorized the bird 
species detected during our surveys into species groups 
based on habitat preferences, diet and foraging sub- 
strate, and nesting substrate. These classifications were 
based on observations reported in the literature (Ehr- 
lich et al. 1988; Robbins et al. 1989a; Freemark and 
Collins 1992; Canterbury et al. 2000); we did not in- 
clude our survey data in these classifications. In addi- 
tion, we created a conservation concern grouping that 
included species which were experiencing statistically 
significant population declines as indexed by North 
American Breeding Bird Survey data for 1966-2000 
(Sauer et al. 2001*). For the purposes of these analyses 
we focused on the groups to which Cerulean Warblers 
belonged: mature forest habitat (n = 13 species), insect- 
foliage foragers (n = 15), canopy_nesters (n = 12), 
and species of concern (n = 14). 

We used two methods to test if the distribution of 
Cerulean Warblers was representative of the distribu- 
tions of other species in the same functional group. 
First, we used the checkerboard score (C-score) devel- 
oped by Stone and Roberts (1990) to test for non- 
randomness in presence-absence matrices. One of the 
reasons we selected this metric of co-occurrence is that 
is not particularly prone to Type 1 error (Gotelli and 
Entsminger 2000*) and, unlike other co-occurrence 
metrics, it allows for overlap in species distributions 
(Gotelli and McCabe 2002; Feeley 2003). We calcu- 
lated C-scores (hereafter, observed C-score) for each 


2004 


JONES, MCLEISH, and ROBERTSON: CERULEAN WARBLER MANAGEMENT 


TABLE |. Interpretation of the principal components axes from analysis of 5 vegetation variables for 59 point-count stations. 


Bold-face eigenvalues significant following bootstrap analyses. 


Axis cvi Ce Wes SPD‘ SSD* LSD! Interpretation of positive axis values 
PCl 29.2 0.6249 -(.6281 -0.2509 0.0187 0.3195 mature forest with dense canopy 
REZ 54.1 0.2676 0.2143 0.7024 0.5205 0.3438 mid-succession forest 

PC3 74.2 0.0036 0.3208 0.1185 -0.7046 0.6218 mid- to late-succession forest 

PC4 88.5 -0.4737 0.0870 -0.4715 0.4648 0.5742 mature forest with patchy canopy 
PES 100.0 0.5599 0.6702 -0.4553 0.1280 = -0.1171 early succession forest 


‘Cumulative variance explained; "Canopy cover (% cover > 12.0 m); ‘Understory cover (% cover < 6.0 m); ‘Sapling density 
(stems/m7); ‘Stem density (stems/m7) 3.0-15.0 cm dbh; ‘Stem density 15.0-30.0 cm dbh. 


functional group to examine if species within each 
functional group were distributed randomly across the 
landscape with respect to one another. For each survey 
year, we used re-sampling techniques to calculate 
10 000 C-scores based on the original presence-absence 
matrix. The observed C-score was then compared with 
the generated distribution of expected C-scores. A C- 
score significantly greater than expected indicates that 
the assemblage is competitively structured; that is, 
individual species have distinct, and often exclusive, 
distributions (Stone and Roberts 1990). Conversely, a 
C-score significantly smaller that expected indicates 
that there is a degree of cohesion in the distribution 
patterns of the species included in the matrix. All C- 
score calculations were carried out using EcoSim 5.0 
(Gotelli and Entsminger 2000). For all iterations, the 
number of species detected at each point-count station 
was kept consistent with the original matrix and each 
station was equally likely to be inhabited by a given 
species. Survey results from 1997 and 1998 were ana- 
lyzed separately. 

In our second test, we used logistic regression ana- 
lyses to predict the probability of occurrence along a 
habitat gradient for all species within each species 
group to which Cerulean Warblers belonged. The 
Red-eyed Vireo (Vireo olivaceus) was not included in 
these analyses due to its near-ubiquitous distribution. 
For these analyses we included only the 1997 surveys 
of the 59 point-count stations for which we collected 
vegetation data. We generated the habitat gradient by 
entering all five vegetation variables into a principal 
components analysis, giving us an approximate 10:1 
ratio of sites to variables. We tested the significance 
of the eigenvalues for each variable within each com- 
ponent using a bootstrap approach, following the 
recommendations of Peres-Neto et al. (2003). We 
drew 1000 bootstrap samples by resampling entire rows 
with replacement, thereby ensuring that the bootstrap 
matrices had the same dimensions as the original veg- 
etation matrix; we conducted a PCA on each of these 
1000 matrices. P-values were estimated by the number 
of bootstrap loadings equal to or less than zero for 
original loadings that were positive (greater than or 
equal to zero for negative loadings), divided by 1000. 


For the purposes of this paper, we considered P = 0.10 
as our significance cut-off. The first PC axis (PC1) 
explained 29% of the variance of the vegetation data 
(Table 1); positive values along PC1 represented sites 
of mature deciduous forest and negative values repre- 
sented early successional forest. We included PC1 as 
the independent variable in our logistic regression ana- 
lyses. All these analyses were performed with JMP 
IN 4.0.2 (SAS Institute Inc. 2000*) and MS-EXCEL 
using the PopTools add-in module (Hood 2002). We 
calculated 95% confidence intervals around the logis- 
tic regression coefficient to facilitate comparison of 
occurrence probabilities across species. 


Results 

(1) Biodiversity indicator — The presence of Ceru- 
lean Warblers was not a significant predictor of avian 
species richness in either 1997 or 1998. In other words, 
observed species richness at Cerulean Warbler point- 
count stations was not significantly different from spe- 
cies richness expected by chance, based on bootstrap 
analyses (1997, observed richness 12.69 + 0.75 spe- 
cies, expected 13.95 + 0.03, P = 0.12; 1998, observed 
9.75 + 0.63, expected 9.82 + 0.03, P = 0.83). 


(2) Umbrella species — All three of the species 
groups to which Cerulean Warblers belong exhibited 
cohesive distributions in both 1997 and 1998, as in- 
dexed by their C-scores (Table 2). C-scores that are 
significantly lower then expected are indicative of co- 
occurrence in the distribution patterns of the species in- 
cluded in the analyses. Species of conservation concern 
also exhibited cohesive distributions in both 1997 and 
1998 (Table 2). 

The results of the logistic regression analyses using 
PC1 indicate that species within each of the species 
groups to which Cerulean Warblers belonged were 
distributed differentially along the generated habitat 
gradient, some more so than others (Table 3). Within 
the mature forest group, only one of the 12 group spe- 
cies (Black-and-white Warbler) did not overlap with 
the Cerulean Warbler confidence intervals. Similarly, 
only one of the 12 canopy nesting species (Blue Jay, 
Cyanocitta cristata) did not overlap with the Cerulean 
Warbler confidence intervals. Conversely, the insect- 


232 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE 2. Tests for cohesive distributions of functional groups, as indexed by the C-score of Stone and Roberts (1990). Esti- 
mated values are mean + SE for a generated random distribution based on the actual presence-absence matrix for each group 
for each year. C-scores that are significantly lower then expected are indicative of co-occurrence in the distribution patterns 


of the species included in the analyses. 


1997 
Functional Group Observed Expected 
Mature forest 80.76 91.94 + 0.03 
Insect-foliage 86.02 92.29 + 0.02 
Canopy nesters 67.03 71.92 + 0.02 
Species of concern 136.11 147.68 + 0.12 


1998 
P Observed Expected ie 
0.0003 41.42 48.13 + 0.02 0.0017 
0.0050 52.96 65.15 + 0.02 <0.0001 
0.0078 36.63 40.90 + 0.02 0.0068 
0.0049 73.32 85.04 + 0.03 0.0001 


TABLE 3. Logistic regression coefficients and upper and lower 95 % confidence intervals (C. I.) predicting occurrence across 
a habitat gradient for members of the mature forest (MF), insect-foliage (IF), canopy nesting (CN) and conservation 
concern (CC) species groups detected in 1997. The confidence intervals of the species in bold face do not overlap with the 


confidence interval of the Cerulean Warbler. 


Species Functional Regression Lower Upper 
group coefficient Spy Vo Osill, 951701 11: 
Cerulean Warbler (Dendroica cerulea) all 0.519 0.121 0.917 
American Redstart (Setophaga ruticilla) all 0.144 -0.299 0.587 
Ovenbird (Seiurus aurocapilla) MF, CC 0.475 0.024 0.926 
Black-and-white Warbler (Mniotilta varia) MEF -0.290 -0.645 0.065 
Scarlet Tanager (Piranga olivacea) all 0.106 -0.239 0.451 
Least Flycatcher (Empidonax minimus) MF, CN, CC 0.116 -0.290 0.522 
Yellow-throated Vireo (Vireo flavifrons) MF, IF, CN 0.066 -0.381 0.513 
Wood Thrush (Hylocichla mustelina) MF, CC -0.287 -0.791 0.217 
Black-throated Green Warbler (Dendroica virens) MF, CN, CC 0.134 -0.272 0.540 
Northern Waterthrush (Seiurus noveboracensis) MF -0.195 -0.777 0.387 
Warbling Vireo (Vireo gilvus) IF -0.995 -1.936 -0.054 
Ruffed Grouse (Bonasa umbellus) MF -0.140 -0.620 0.340 
Veery (Catharus fuscescens) MF, CC -0.622 -1.786 0.542 
Yellow-rumped Warbler (Dendroica coronata) MF, IF, CN 0.009 -0.483 0.501 
Common Yellowthroat (Geothlypis trichas) IEXEE -0.192 -0.555 0.171 
Golden-winged Warbler (Vermivora chrysoptera) IEVEGE -0.534 -0.955 -0.113 
Indigo Bunting (Passerina cyanea) TENCE -0.535 -0.974 -0.096 
Chestnut-sided Warbler (Dendroica pensylvanica) IEF; Ee -0.705 -1.438 0.028 
Yellow Warbler (Dendroica petechia) IF -0.232 -0.585 0.121 
Black-capped Chickadee (Poecile atricapillus) IF 0.058 -0.279 0.395 
Rose-breasted Grosbeak (Pheucticus ludovicianus) Wee -0.725 -1.270 -0.180 
Baltimore Oriole (Icterus galbula) LE ENEC 0.281 -0.217 0.779 
Blue-grey Gnatcatcher (Polioptila caerulea) IF, CN -0.279 -0.781 0.223 
Cedar Waxwing (Bombycilla cedrorum) CN -0.066 -0.440 0.308 
Blue Jay (Cyanocitta cristata) CN -0.873 -1.528 -0.218 
Eastern Wood-Pewee (Contopus virens) CN, CC -0.202 -0.670 0.266 
American Crow (Corvus brachyrhynchos) CN -0.304 -0.835 0.227 


foliage and conservation concern groups exhibited less 
concordance with the Cerulean Warbler distribution 
relative to the habitat gradient — 43% (6 of 14) and 
29% (4 of 14), respectively. 


Discussion 

Effective biodiversity indicators tend to be habitat 
specialists with wide geographic ranges; they also tend 
to have well-known natural histories (Caro and O’ Do- 
herty 1999). Despite fitting this profile (Hamel 2000; 
Oliarnyk 1996; Jones 2000; Barg 2002), our results 
indicate that the Cerulean Warbler would not be par- 


ticularly effective as an avian biodiversity indicator in 
eastern Ontario, as its distribution across the studied 
landscape did not coincide with areas of high avian 
species richness. 

Our co-occurrence and logistic regression results do 
suggest that the Cerulean Warbler is suited to a role as 
an umbrella species. Perhaps not surprisingly, the dis- 
tribution of the Cerulean Warbler was well matched 
to the distributions of other canopy nesters in mature 
deciduous forest (e.g., Scarlet Tanager, Piranga oli- 
vacea). Given the hierarchical nature of habitat selec- 
tion, broad habitat requirements are likely more impor- 


2004 


tant in determining species distributions than are spe- 
cies’ food and nesting requirements (Hutto 1985; Block 
and Brennan 1993); indeed, we found that the species 
grouped by diet were more variable in their distri- 
butions along our succession gradient than were the 
species grouped by habitat type or nest location. Small 
body size — and, as a consequence, small home range 
size (Barg et al. in press) — could limit the Cerulean 
Warbler’s umbrella suitability, as effective umbrella 
species tend to have large home ranges (Caro and 
O’Doherty 1999); however, the Cerulean Warbler’s 
apparent tendency to live in conspecific aggregations 
(Hamel 2000) potentially offsets any limitation im- 
posed by small body size and individual home range 
size. We do not mean to suggest that the Cerulean 
Warbler would be a better umbrella than other, more 
widespread, species such as the Scarlet Tanager. Rath- 
er, our conclusions suggest that, if habitat manage- 
ment is directed at Cerulean Warblers, other species 
will likely benefit. 

Our co-occurrence and logistic regression results 
highlight a potential management conflict between 
mature forest and shrubland species. Although our C- 
scores indicate significant patterns of co-occurrence 
within the conservation concern group, there was no 
overlap between the logistic regression confidence in- 
tervals of the Cerulean Warbler and shrubland species 
that are considered to be at risk, such as the Golden- 
winged Warbler (Vermivora chrysoptera). Given their 
disparate habitat requirements (i.e., forest edge vs. 
forest interior), management for Golden-winged War- 
blers will necessarily conflict with management aimed 
at maximizing Cerulean Warbler population health. 

One additional conservation role the Cerulean War- 
bler may fill, and may be already filling, is that of a 
flagship species: a species that attracts attention sup- 
port by virtue of its ‘charismatic’ nature (Simberloff 
1998). The Cerulean Warbler is a Neotropical migrant 
songbird, a group of birds that has been in the conser- 
vation spotlight over the last 30 years (e.g., Robbins 
et al. 1989b). The Cerulean Warbler is a beautiful 
bird with an elusive nature that has a high profile due 
to extensive public education and activism (Rosenberg 
et al. 2002) and is highly valued by birders and orni- 
thologists alike. More recently, the Cerulean Warbler 
has achieved notoriety as the focus of a legal battle 
surrounding its candidacy for listing on the U.S. En- 
dangered Species Act (Ruley 2000). This attention 
has led to the development of the CWTG and has 
spurred a great deal of basic research and forest man- 
agement interest (Hamel et al. 2004). The apparent 
dependence of this species on large tracts of forest 
creates a possibility for the Cerulean Warbler to serve 
as a valuable symbol of the overall health of deciduous 
forests in eastern North America. 


Acknowledgments 
We thank J. Pither and R. Montgomerie for sta- 
tistical advice. J. Barg, A. Erskine and an anonymous 


JONES, MCLEISH, and ROBERTSON: CERULEAN WARBLER MANAGEMENT 


reviewer provided helpful comments on earlier drafts. 
F. Phelan, M. Phelan, F. Connor and the staff at the 
Queen’s University Biological Station provided logis- 
tical support. Funding was provided by B. and K. 
Jones, Wildlife Habitat Canada, the Eastern Ontario 
Model Forest Program and an NSERC operating 
grant to RJR. We also had the support of the World 
Wildlife Fund of Canada in the form of a MacNaughton 
Conservation Scholarship to JJ and grants from the 
Endangered Species Recovery Fund to RJR. This 
project is part of Natural Legacy 2000, a nationwide 
initiative in Canada to conserve wildlife and habitats 
in private and public. We gratefully acknowledge the 
support of the Government of Canada’s Millennium 
Partnership Fund. JJ was also supported by an NSERC 
scholarship, Queen’s Graduate Awards and the Queen’s 
Bracken Fellowship. 


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Gotelli, N. J., and G. L. Entsminger. 2000. EcoSim: null 
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Hood, G. 2002. PopTools, Version 2.3. http://www.cse. 
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Mathsoft, Inc. 1997. S-Plus 4.0. Data Analysis Products 
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Rosenberg, K. V., S. E. Barker, and R. W. Rohrbaugh. 
2002. An atlas of Cerulean Warbler populations — Final 
report to United States Fish and Wildlife Service: 1997- 
2000 breeding seasons. Cornell Lab of Ornithology, Ithaca, 
New York. 

Ruley, D. A. 2000. Petition under the Endangered Species 
Act to List the Cerulean Warbler, Dendroica cerulea, as a 
Threatened Species. Southern Environmental Law Center, 
Asheville, North Carolina. 

SAS Institute Inc. 2000. JMP IN 4.0.2. SAS Institute Inc., 
Cary, North Carolina. 

Sauer, J. R., J. E. Hines, and F. Fallon. 2001. The North 
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Received 22 October 2002 
Accepted 21 July 2004 


Parasite Prevalence in Dark-eyed Juncos, Junco hyemalis, Breeding at 
Different Elevations 


HEATHER BEARS 


Department of Zoology, University of British Columbia, 6270 University Boulevard, Vancouver, British Columbia V6T 174 
Canada; e-mail: bears @zoology.ubc.ca 

Present address: Centre for Applied Conservation Research, 2424 Main Mall, Forest Sciences Building, University of 
British Columbia, Vancouver, British Columbia V6T 1Z4 Canada 


Bears, Heather. 2004. Parasite prevalence in Dark-eyed Juncos, Junco hyemalis, breeding at different elevations. Canadian 
Field-Naturalist 118(2): 235-238. 


During the summer of 2001, Dark-eyed Juncos (Junco hyemalis) were captured within the lowest (1000 m above sea level) 
and highest (2000 m asl) elevation extremes of their breeding range in Jasper National Park, Alberta. Blood samples were 
taken to identify parasite genera, and to test for differences in parasite prevalence among elevations. The most common 
parasites at either elevation were Haemoproteus spp., Leucocytozoon spp., and Trypanosoma spp. A significantly higher 
proportion of low- compared to high-elevation birds was infected by at least one of these, supporting the prediction that 


high-elevation habitats may be refuges from parasites. 


Key Words: Dark-eyed Juncos, Junco hyemalis, blood parasites, mountains, elevation, Alberta. 


Birds breeding at high elevations often experience 
a delayed date of reproductive onset, exposure to ex- 
treme weather, and scarce or sparsely distributed re- 
sources (Hamman et al. 1989; Landmann and Winding 
1993; Kollinsky and Landmann 1996; Widmer 1999; 
Bears et al. 2003). Dark-eyed Juncos (Junco hyemalis) 
show a significant reduction in seasonal reproductive 
output with increasing breeding elevation from the 
montane valley (1000 m asl) to the subalpine-alpine 
treeline (2000 m asl) in Jasper National Park, Alberta 
(Bears 2002, Bears et al. 2003). Yet, high elevation 
habitats are not occupied by less competitive age or 
size classes of juncos, or by later arriving individuals 
(Bears 2002), all of which are traits that render juncos 
less successful competitors for territory (Cristol et al. 
1990; Grasso et al. 1996). Further, inter-annual return 
of birds to their site of capture is high, and roughly 
equivalent among elevations (Bears 2002). Hence, it 
does not appear that juncos are “forced” to breed at 
high elevations due to intraspecific competitive exclu- 
sion from lower-elevation habitat. This led me to ex- 
plore the idea that high-elevation juncos are compen- 
sated for decreased seasonal reproductive output over 
their lifetimes by some unrecognized benefits of breed- 
ing at high elevations. In other mountain ranges, 
studies have shown that the blood parasites of birds, 
and the insect vectors that carry them, tend to be low 
at upper breeding elevations (Stabler et al. 1974; Braun 
et al. 1993). Thus, as part of a series of investigations 
into the benefits of high-elevation breeding, I tested 
whether the juncos have fewer parasites within higher- 
versus lower-elevation segments of their breeding 
range in Jasper. 


Methods 

Juncos were captured and monitored at eight 50 — 
70 ha study sites in Jasper National Park (52°53'N, 
118°3'W), Alberta, from 1 May to 20 August in 2000, 
and from 15 April to 20 August in 2001. Four sites 
were located near the lowest elevation within the Park 
(1000 — 1020 m asl), separated by 5-10 km. Four sites 
were at the highest elevation at which juncos breed 
within the park (1950-2100 m asl), and were separat- 
ed by > 18 km. All high-elevation sites had south or 
southest aspects. Morphological measurements taken 
from juncos in these sites (H. Bears, unpublished data) 
most closely match the Oregon subspecies, Junco 
hyemalis oregonus (Miller 1941), but some appear to 
be intergrades between Junco hyemalis oregonus and 
Junco hyemalis hyemalis, which produces Junco hye- 
malis cismontanus (Miller 1941). Subspecies desig- 
nations of Dark-eyed Juncos are a capricious topic 
based on phenetic rather than genetic data, and they 
are a taxonomic nightmare in the Canadian Rockies 
where multiple subspecies meet and hybridize. There- 
fore, we refer to the birds analyzed here simply as 
Junco hyemalis. 

Birds were captured using Japanese mist nets with 
painted model male juncos as decoys. Juncos were 
lured towards the net by playing the taped song of a 
conspecific. Birds were given a numbered Canadian 
Wildlife Service leg band and colour bands that 
conveyed sex, age, and site information when caught. 
Blood samples were taken from each bird caught 
between | May and 20 August in 2001 only by punc- 
turing the alar vein with a needle and collecting ca. 
40 ul of blood in a heparinized microhematocrit tube. 


235 


Blood was blown out of microhematocrit tubes into 
centrifuge tubes and kept on wet ice. Because blood 
parasites may follow a diurnal periodicity (Gore et 
al. 1982), the birds selected for parasite identification 
were caught at approximately the same time each day 
(7:00-12:00 hrs). Within 10 hours of collection, 5 ul 
of blood were used to produce three microscope slides 
per bird, using the methods of Harrison and Harrison 
(1986) and Bennett (1970). The slides were fixed in 
100% methanol immediately. In the laboratory, blood 
smears were stained with Geimsa stain for 30 min- 
utes, and rinsed with distilled water followed by acetone 
under a fumehood (Deviche et al. 2001). Smears were 
examined under 400 x 10 magnification, and 50 fields 
of view per slide were classified as negative or posi- 
tive for various parasites. Parasites were identified to 
the genus level by examination under high power 
(1000 x magnification) with oil immersion, using 
various keys (Pierce 1981; Bennett and Pierce 1988; 
Burrey-Caines and Bennett 1992; Bennett et al. 1994), 
~and by comparing with photos of blood parasites taken 
from juncos captured in Alaska (supplied by Pierre 
Deviche, Arizona State University, personal commu- 
nication). Parasite species were difficult to ascertain 
with absolute certainty, but the genus level could be 
resolved without ambiguity. Therefore, we used the 
genus level when comparing parasite prevalence among 
elevations. 

At each site, one bird was caught between the 1* and 
5" of each month (May-August), and another between 
the 11 and 15" of each month, in order to represent 
sites and time periods within the breeding season equiv- 
alently. No females were captured using playback 
and mist-netting methods, and so analyses here deal 
solely with males. The statistical significance level 
for all tests conducted was set at a = 0.05. All tests 
were performed using SPSS 10. All techniques used 
were approved by the animal care committee of the 
University of British Columbia (A0-0046), Parks Can- 
ada (2000-008), and Environment Canada (Banding: 
10429 AJ; Collection: BC SCI 2000/067). 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Results 

Thirty-two high- and 32 low-elevation adult male 
birds caught across all eight sites were used in these 
analyses. In addition, 12 fledglings were caught at high 
elevations (in August) and 12 were caught at low ele- 
vations (in August). Three parasite genera were found 
in the blood of juncos: Haemoproteus (Family Plas- 
modiidae), Leucocytozoon (Family Plasmodiidae) and- 
Trypanosoma (Family Trypanosomatidae). All data are 
summarized in Table 1. The most prevalent blood para- 
site at both elevations was Haemoproteus, followed 
by Leucocytozoon, and Trypanosoma. A significantly 
higher proportion of low-elevation birds (66%, 21/32) 
compared to high-elevation birds (28%, 9/32) were 
infected by at least one of these parasites (P = 0.002, 
Fisher’s Exact Test). A higher percentage of low-ele- 
vation birds (34%, 11/32) compared to high-elevation 
birds (19%, 6/32) were infected with Haemoproteus 
spp., but the difference was not significant (P = 0.08, 
Fisher’s Exact Test). Similarly, more low-elevation 
birds (25%, 8/32) versus high-elevation birds (9%, 
3/32) were infected with Leucocytozoon spp., with a 
near significant difference (P = 0.07, Fisher’s Exact 
Test). Of these individuals, 42% (8/19) of low-eleva- 
tion and 44% (4/9) of high-elevation birds had both 
Haemoproteus spp. and Leucocytozoon spp. present 
in their blood. Two cases of infection with Trypano- 
soma spp. were found in low-elevation birds, and none 
at high elevations. Thirty-four percent (11/32) of low- 
elevation and 72% (23/32) of high-elevation birds 
were not infected with any parasites. There was no 
relationship between date of capture and infection at 
low elevations (P = 0.23, r = 0.33; Pearson’s r). How- 
ever, at high elevations, a low proportion of individ- 
uals (12.5%, 2/16) caught between 1 May and 15 June 
were infected with Leucocytozoon spp. and Haemo- 
proteus spp., but between 15 June and 30 August, a 
significantly higher proportion were infected (7/16, 
44%; P = 0.05, Fisher’s Exact Test). In samples from 
low elevations not used in this analysis (taken from 
birds nearby, but not within our study areas) two cases 


TABLE |. Summary of data on presence and absence of blood parasites in Dark-eyed Juncos from low (~1000 m asl) and 


high (~2000 m asl) elevations. 


Adults (ASY) Infection/Parasite 


Infected 

Trypanosoma present 
Leucocytozoon present 
Haemoproteus present 
Both Haemoproteus and 
Leucocytozoon present 
no parasites 


Fledglings Parasite 


Leucocytozoon present 


Low (N = 32) High (N = 32) 
21 9 
2 0 
8 3 
11 6 
8 of 19 individuals 4 of 9 individuals 
11 23 
Low (N = 12) High (N = 12) 
2 1 


2004 


of Plasmodium spp. (family Plasmodiae) were noted. 
More fledglings were infected with Leucocytozoon spp. 
at low elevations as compared to at high elevations 
(17%, 2/12, vs 8.3%, 1/12), but the difference was not 
significant (P = 0.39, Fisher’s Exact Test). No other 
blood parasites were observed in fledglings. 


Discussion 

High-elevation adult males and fledglings had lower 
incidences of blood parasites as compared to low-ele- 
vation adult males and fledglings. If relief from para- 
sites enables adults to survive longer and breed for 
more years, then this could aid in equalizing the life- 
time reproductive success of high- and low-elevation 
birds. Blood parasites can decrease survivorship by 
directly increasing susceptibility to predation (Vaughn 
and Coble 1975). In addition, there may be a trade- 
off between reproductive effort and the efficiency of 
the immune response. For instance, parasite loads may 
increase during the reproductive period, as they did 
here at high elevations, when breeding adults spend 
considerable time provisioning their young and in 
nest and territorial defence, or when breeding effort 
is increased experimentally (Ots and Horak 1996; 
Norris et al. 1994; Weatherhead and Bennett 1991, 
1992; Rintmaki et al. 1999). A simpler reason for the 
differential susceptibility observed here might also be 
due to a difference in the timing of emergence of the 
insect vectors carrying the parasites. 

The two most common parasite genera in this study, 
Haemoproteus and Leucocytozoon, are protozoon para- 
sites of birds. Haemoproteus spp. are primarily trans- 
mitted by insects in the dipteran family Ceratopogoni- 
dae (no-see-ums, sandflies) or Hippoboscidae (louse 
flies), whereas Leucocytozoon spp. are primarily trans- 
mitted by simuliids (e.g., black flies) (Greiner and 
Ritchie et al. 1994; Rosskopf and Woerpel 1996; Rint- 
maki et al. 1999). Trypanosoma (likely avium), which 
was rare in our study, is also transmitted by members 
of the family Simuliidae. The principal effects of 
Leucocytozoon infections are intravascular haemolytic 
anemia, weight loss, and sometimes death, whereas 
Haemoproteus and Trypanosoma in birds are generally 
less pathogenic (Greiner and Ritchie 1994). Finally, 
two low-elevation samples (not analyzed as part of the 
sub-samples selected) contained Plasmodium spp., 
which is transmitted by mosquitos (family Culicidae). 
Almost all of these insect vectors were noted to have 
emerged later at high elevations, perhaps preventing 
exposure of birds to the parasites for much of the sea- 
son. High-elevation birds were also at lower densities 
(Bears 2002), and therefore transmission between 
birds may have been lower. 

Results suggest that high-elevation habitats may be 
of conservation importance in limiting the spread of 
blood parasites in birds. The Canadian Rockies may 
play a particularly important role in limiting the rate 
at which avian diseases travel east and west across 


BEARS: PARASITE PREVALENCE IN DARK-EYED JUNCOS 


the Rocky Mountains. Further work, including com- 
parisons of parasite levels in females, and in other 
species that breed over wide elevation ranges, is re- 
quired in order to assess the generality of this parasite 
refugium hypothesis in the Rocky Mountains of Can- 
ada. Mechanisms by which high-elevation birds are 
protected from parasites should also be explored. 


Acknowledgments 

Project funding was provided by NSERC, and 
personal funding by NSERC PGS-A, Environment 
Canada, Agriculture and Agri-foods Canada, Health 
Canada, Fisheries and Oceans Canada, and Natural 
Resources Canada. J. N. M. Smith provided helpful 
advice in the preparation of this manuscript. S. Stearns 
encouraged me with the pursuit of parasite quantifi- 
cation across elevations. I benefited in the field from 
the help of G. Brown, M. Bandura, and A. Bendzsak. 
M. Adamson provided microscopy facilities. Thanks 
to A. J. Erskine and two anonymous referees for 
reviewing the manuscript. 


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Received 24 October 2002 
Accepted 4 November 2004 


Movements of Subadult Male Grizzly Bears, Ursus arctos, in the 
Central Canadian Arctic 


RosBert J. GAu!, Puitip D. MCLOUGHLIN, RAY Case!, H. DEAN CLUFF?, ROBERT MULDERS!, and 
FRANCOIS MESSIER? 


'Wildlife and Fisheries Division, Department of Resources, Wildlife and Economic Development, Government of the Northwest 
Territories, #600 5102-50th Avenue, Yellowknife, Northwest Territories X1A 3S8 Canada; e-mail: rob_gau@ gov.nt.ca 

*Department of Biology, University of Saskatchewan, 112 Science Place, Saskatoon, Saskatchewan S7N 5E2 Canada 

3Department of Resources, Wildlife, and Economic Development, Government of the Northwest Territories, North Slave 
Region, P.O. Box 2668, Yellowknife, Northwest Territories X1A 2P9 Canada 


Gau, Robert J., Philip D. McLoughlin, Ray Case, H. Dean Cluff, Robert Mulders, and Francois Messier. 2004. Movements of 
subadult male Grizzly Bears, Ursus arctos, in the central Canadian Arctic. Canadian Field-Naturalist 118(2): 239-242. 


Between May 1995 and June 1999, we equipped eight subadult male (3-5 yrs old) Grizzly Bears (Ursus arctos) with satellite 
radio-collars within a study area of 235 000 km?, centred 400 km northeast of Yellowknife, Northwest Territories, Canada. 
Subadult male annual home ranges were extraordinarily large (average = 11 407 km’, SE = 3849) due, in part, to their 
movement’s occasional linear directionality. We believe their long-range linear movements may reflect some individuals 
tracking the migration of Caribou (Rangifer tarandus). Seasonal daily movement patterns were similar to adult males that 
were previously reported. The areas used by these bears are the largest ranges reported for any Grizzly Bears and the scale 
of their movements may put individual bears in contact with humans even when developments are hundreds of kilometres 


from the central home range of an animal. 


Key Words: Grizzly Bear, Ursus arctos, home range, movements, subadult, central Arctic, Northwest Territories. 


The population of Barren-ground Grizzly Bears 
(Ursus arctos) in the Northwest Territories (NWT) 
exists at a low density in the tundra and is considered 
“sensitive” after a recent species-at-risk assessment 
(Government of the NWT, 2000). As such, there is 
concern for any increase in human presence that might 
impact on their numbers. 

Economic activity in the central Arctic of the NWT 
and Nunavut increased dramatically in the early 1990s 
with the discovery of diamonds. At present there are 
three mines (two diamond, one gold) and numerous 
base-metal, gold, and other diamond developments in 
the region. To address the effects of these develop- 
ments, in 1995 the Government of the NWT and the 
University of Saskatchewan initiated the first multi- 
faceted research program into the ecology of Barren- 
ground Grizzly Bears inhabiting the central Arctic. 

One aspect of our research program was to detail 
the spatial requirements of the Barren-ground Grizzly 
Bears in this region (McLoughlin 2000). However, 
incidental to our primary objectives, we obtained data 
on large-scale movements of some subadult male bears 
(3-5 yrs old) that were also captured and monitored. 
Although many aspects of bear ecology have been well 
documented, descriptions of dispersal and movement 
patterns for subadult Grizzly Bears remain rare and 
difficult to obtain (McLellan and Hovey 2001). While 
progress with DNA analysis may shed insights on sub- 
adult dispersal in the future (see Waits et al. 1999; 
Woods et al. 1999; Woods and Strobeck 2000), current 
methods to examine dispersal in bears generally re- 


quire marking young animals to identify when captured 
or killed at a later date, or radio tracking 2- and 3-year 
olds captured prior to or immediately following sepa- 
ration from their mother. Our findings parallel and 
complement McLoughlin et al. (1999) and McLoughlin 
et al. (2003). 


Methods 

The study area was centred in Canada’s central 
Arctic (66°10'N, 111°25'W), encompassing approxi- 
mately 235 000 km? of mainland Nunavut and the 
NWT (Figure 1). The study area was delineated by the 
community of Kugluktuk, the Kent Peninsula, Aylmer 
Lake, Mackay Lake, and Great Bear Lake. We previ- 
ously noted the biophysical characteristics of the region 
in Gau et al. (2002). 

Between May 1995 and June 1999, helicopters were 
used to search for and capture bears. Bears weighing 
>110 kg (males) and >90 kg (females) were fitted with 
tracking devices. There were the minimum weights we 
considered bears robust enough to wear a collar safely. 
We considered most two- and three-year-old bears too 
small and growing too rapidly to be fitted with satel- 
lite collars. Some telemetry methods (e.g., inserts or 
breakaways to collar belting, ear-tag transmitters, ex- 
pandable radio-collars) show promise; however, im- 
provements are still needed (Costello et al. 2001). 

Satellite (Service Argos Inc., Landover, Maryland, 
USA) and conventional VHF radio-telemetry (Telonics 
Ltd., Mesa, Arizona, USA) were used to obtain spatial 
information on Barren-ground Grizzly Bears. Most 


239 


240 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE 1. Subadult male Grizzly Bears, and their periods of long-range (>200 km) linear directional movement, captured 
and collared between 1995 and 1999 in the central Canadian Arctic. 


Number 95% fixed Mean Daily Movement (km/day) Linear Directional Movements 
of kernal Late Distance Number 
Bear Age Year locations range (km?) Spring Summer summer Autumn covered (km) of days 
G595 4 1995 103 15 899 eS) 10.4 6.2 9) 539 Sif) 
G600 3 1995 129 22 007 16.6 8.8 10.5 es 7719 59 
G612 4 1995 91 4448 24.8 9.7 4.0 Nile ge 471 23 
446 33 
G618 5 1995 66 4540 6.1 6.7 9.5 5.6 
G656 5) 1996 45 32 188 n/a! n/a! n/a! n/a! 273 26 
305 40 
G657 5) 1996 42 6162 n/a! n/a! n/a! n/a! 
G689 4 1998 58 3662 n/a! n/a! n/a! n/a! 
G700 4 1998 64 2349 7.9 59 4.2 ay 201 32 


' <8 locations/season in every season of the year were recorded thus omitted from analysis. 


collars were designed to transmit approximately 2-5 
latitude-longitude locations every two days (8-hour 
duty cycle) from 1 May to | November. 

Our calculations and techniques for study of ani- 
mals, ranges from satellite telemetry locations and rates 
of movement (km/day) were previously described in 
McLoughlin et al. (1999) and McLoughlin et al. (2003). 
Annual home ranges were determined using the 95% 
isopleth for bears only with 238 locations, so as not 
to overestimate range size with smaller sample sizes 
(Seaman et al. 1999). Also, only those animals that 
transmitted 28 locations/season in every season of the 
year were included for analysis. We defined seasons 
according to changes in the diet of Barren-ground 
Grizzly Bears during the active period (adapted from 
Gau et al. 2002), including spring (den emergence- 
20 June), summer (21 June-31 July), late-summer (1 
August-9 September), and autumn (10 September- 
den entrance). 


Results 

Of the male bears that were not in a family group 
with their mother, 6 of the 45 males handled were too 
small for collaring. However, we did collar and mon- 
itor 8 subadult males between 3 and 5 years of age 
out of a sample of 39 collared males (Table 1). Male 
subadult movements were extraordinarily large and 
annual home range averaged 11 407 km? (SE = 3849, 
range 2349 — 32 188). 

A unique feature of the subadult movements we 
observed consisted of periods of long-range (>200 km) 
linear directional forays. Five of the eight subadults 
we followed exhibited this linear movement pattern. 
For example, 4-year-old bear G612 in 1995 moved 
471 km from 31 May to 22 June along a northerly 
trek from the treeline to the Arctic coast. While the 
other movements may not be as dramatic as G612 in 
terms of time a distance was covered, long-range linear 
directional movement appeared common for subadult 


males in this Barren-ground Grizzly population. Bears 
G595, G600, G612, and G656 had linear directional 
movements in the spring; additionally, bears G612, 
G656, and G700 had autumn linear directional move- 
ments. 

The subadult males followed a seasonal daily move- 
ment pattern similar to adult males (see McLoughlin 
et al. 1999). Only five bears met our criteria to inves- 
tigate seasonal movements. Means were 12.6 km/d 
(SE='3.6), 8:3 km/d' (SE = 0:9), 6:9’ kin/d"\(SE'=' 1:3); 
and 7.6 km/d (SE = 1.2) for spring, summer, late 
summer, and autumn, respectively. There was a clear 
general trend from a high rate of movement (spring) to 
lower rates for the rest of their active period, although 
an ANOVA determined no significant differences 
between seasonal means (F3 ,,= 1.6, P = 0.2). 


Discussion 

The annual ranges of adult and subadult Barren- 
ground Grizzly Bears in the central Canadian Arctic 
are the largest ranges yet reported for grizzlies in North 
America (see Table 1, McLoughlin et al. 1999). How- 
ever, there are very few published data about subadult 
Grizzly Bear movement patterns either to refute or sup- 
port our claim (McLellan and Hovey 2001). Jonkel 
(1987), along with Mace and Waller (1997), noted 
that dispersing immature brown bears tended to have 
smaller home ranges than adult male bears. However, 
LeFranc et al. (1987), along with Nagy et al. (1983), 
summarized examples where subadults, believed dis- 
persing from maternal home ranges, had home ranges 
as large or larger than those of adult males. Regardless, 
the magnitude of the movement patterns we observed 
eclipses other results that have been previously pub- 
lished. 

We have gleaned valuable insights into the move- 
ment patterns of Canadian Barren-ground Grizzly 
Bears. Mean annual ranges were 7245 km? for adult 
males and 2100 km? for females, with no difference in 


2004 


GAU, MCLOUGHLIN, CASE, CLUFF, MULDERS, and MESSIER: MOVEMENTS OF BEARS 241 


Kent Peninsula 


‘CORONATION 
GULF 


Kugluktuk 


hie de oa 


i \ MacKay axe 
a Aylmer 


=z) 
250 km 


co Nunavut 
Northwest 7 


Erritories 


Lake 


FiGureE |. Location of the study area in Canada’s central Arctic. The treeline indicates the approximate northern limit of 


coniferous forest in the region. 


the ranges for females of differing family status (Mc- 
Loughlin et al. 2003). The large disparity between 
adult males and females (5145 km7?), and especially 
between subadult males and females (9307 km2), is in- 
teresting since females introduce younger male bears 
only to a seemingly small portion of the land they will 
eventually use in the central Arctic. Although we can 
expect a high degree of movement and population over- 
lap among males in the central Arctic (McLoughlin et 
al. 2002), it is possible that some of the subadult male 
ranges we recorded were somewhat inflated. We sus- 
pect that some bears in this study tracked the spring 
migration of Caribou (Rangifer tarandus), a behaviour 
previously suspected in northern Alaska for Barren- 
ground Grizzlies (Reynolds and Garner 1987). 

At the daily and seasonal movement rates we ob- 
served for subadult males, because of their similarity to 
adult male daily movement rates, it would be possible 
for subadult males to have home range sizes typical 
of male adults in the region. However, the unique fea- 
ture of the subadult movements we observed appeared 


to be their occasional linear directionality. Other than 
homing behaviours of transplanted bears (Miller and 
Ballard 1982), extended directional movements are 
rare (McLellan and Hovey 2001). Explanations for long- 
range bear movements include seeking out quality hab- 
itat, spacing behaviour resulting from social interac- 
tions with other bears, the abundance and distribution 
of food (e.g., following migrating caribou herds), in- 
breeding avoidance, and maximizing reproductive fit- 
ness, or dispersal immediately following separation 
from their mother (Rogers 1987; Pasitschniak-Arts 
and Messier 2000). All these factors likely contributed 
to the movement patterns of subadult males in the 
Canadian central Arctic. McLoughlin et al. (1999) also 
pointed out that Barren-ground Grizzly Bears have 
larger scale movements when compared to other griz- 
zly populations principally due to low primary pro- 
ductivity in the tundra environment. 

The spatial behaviour of subadult male grizzlies 
increases their probability of coming in contact with 
humans even when sites of human activity (e.g., ex- 


242 


ploration and hunting camps, industrial developments, 
and communities) are of considerable distance from 
the central home range of an individual. Subadult bears 
in particular do not have the life-experience of mature 
bears, are often imbued with more curiosity, and thus 
are highly susceptible to human activity. Management 
of bears in the central Arctic should focus on main- 
taining low levels of human-caused mortality of bears, 
with the realization that communities, hunting camps, 
and mining/exploration camps may impact bears from 
more than just the general vicinity. Also, the wide- 
spread movements of subadult bears are a compelling 
argument for a comprehensive and consistent bear 
conflict avoidance program throughout the central 
Arctic. Developments that might not be considered in 
optimum Grizzly Bear habitat should have the tools 
or resources to effectively manage potential problem 
bears. 


Acknowledgments 

Funding for our research was supplied by the 
Government of the NWT Department of Resources, 
Wildlife, and Economic Development, University of 
Saskatchewan, West Kitikmeot/Slave Society, Indian 
and Northern Affairs Canada, BHP Diamonds, Rescan 
Environmental Services, Axys Environmental Consult- 
ing Ltd., Diavik Diamond Project, Monopros Limited, 
Echo Bay Mines Ltd., Nunavut Wildlife Management 
Board, the Federal Department of Indian Affairs and 
Northern Development, the Northern Scientific Train- 
ing Program, the Natural Sciences and Engineering 
Research Council of Canada, and the Polar Conti- 
nental Shelf Project (number 01298). 


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Gau, R. J., R. Case, D. F. Penner, and P. D. McLoughlin. 
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Received 3 February 2002 
Accepted 11 November 2004 


Severe Chronic Neck Injury Caused by a Snare in a Coyote, Canis latrans 


PIERRE-YVES Daoust!:3 and PETER H. NICHOLSON! 2 


‘Canadian Cooperative Wildlife Health Centre, Department of Pathology and Microbiology (Daoust), and Class of 2002 
(Nicholson), Atlantic Veterinary College, University of Prince Edward Island, 550 University Avenue, Charlottetown, 
Prince Edward Island CIA 4P3, Canada 

Present address: College Village Animal Clinic, 2036 E. Northern Lights Boulevard, Anchorage, Alaska, 99508, USA 

3Corresponding author: e-mail: daoust @upei.ca 


Daoust, Pierre-Yves, and Peter H. Nicholson. 2004. Severe chronic neck injury caused by a snare in a Coyote, Canis latrans. 
Canadian Field-Naturalist 118(2): 243-246. 


A two-year-old male Coyote, Canis latrans, in poor body condition was found in a moribund state with a snare deeply embedded 
in the ventral portion of its neck, more than a month after the official end of the trapping season on Prince Edward Island. 
This snare had presumably malfunctioned, and the cable had cut through the soft tissues of the neck as well as the trachea 
and had obstructed both jugular veins and both common carotid arteries but had largely spared both vagosympathetic trunks. 
Cases like this illustrate the need to continue to work on improving the efficiency of trapping methods, through research and 


trapper education. 


Key Words: Coyote, Canis latrans, trapping, injury, snare, Prince Edward Island. 


Animal welfare issues surrounding trapping meth- 
ods used to capture furbearers have been an ongoing 
topic of discussion between wildlife managers and 
animal protection groups for many years (Proulx and 
Barrett 1991). Because of this and of the economic 
importance of the fur trade to Canada, this country 
has taken a leading role in developing national and 
international standards for the performance of restrain- 
ing- and killing-type traps. In the section of the Agree- 
ment on International Humane Trapping Standards 
(AIHTS) (1997*) dealing with killing type traps such 
as the commonly used rotating-jaw Conibear trap, the 
designated time limit to irreversible loss of corneal 
reflexes in at least 80% of the animals of the target 
species caught in the trap varies from 45 seconds in 
Ermine (Mustela erminea) to 120 seconds in Pine Mar- 
ten (Martes americana) to 300 seconds in 17 other 
species of North American and European furbearing 
animals. This Agreement requires that the parties in- 
volved continue research with a view to lowering the 
threshold requirements. Nonetheless, it implicitly rec- 
ognizes that no killing trapping method used to cap- 
ture wild animals can as yet guarantee a humanely 
acceptable process in each instance. Manual neck 
snares (where the animal provides the energy neces- 
sary to tighten the snare) are widely used by trappers 
to capture various furbearing animals, and Guthery and 
Beasom (1978) and Boddicker (1982) discussed the 
advantages and disadvantages of snares as compared 
to other trapping devices. Snares are allowed in most 
Canadian jurisdictions, except the southern regions 
of some provinces. Being considered devices that are 
constructed by the trappers themselves, they are not 
subject to the testing requirements under the AIHTS 
(1997), but their design must be approved by relevant 
competent authorities such as provincial Departments 


of Wildlife. This article describes an unsuccessful cap- 
ture by snare of a Coyote (Canis latrans), with severe 
consequences. 


Case Description 

On 26 February, 2002, a male Coyote, subsequently 
determined to be 2 years old (Matson’s Laboratory, 
Milltown, Montana, USA), was found alive but mori- 
bund in a field in Kings County (46°08'N, 62°36'W), 
Prince Edward Island (PEI), and was euthanized by a 
shot to the head. It was submitted for necropsy to the 
Diagnostic Laboratory of the Atlantic Veterinary Col- 
lege, University of PEI. Postmortem examination 
revealed that the animal had been caught in a snare 
and had managed to escape. The snare’s cable, made of 
galvanized steel wire 2 mm in diameter, was equipped 
with a standard U-shaped lock, similar to an Adams 
lock. The snare surrounded the animal’s neck, with a 
portion of cable approximately 20 cm long dangling 
beyond the neck. The coyote was in poor body con- 
dition; the stomach was completely empty, and the 
intestines contained only a small amount of ingesta. 
However, some fat was still present around the base 
of the heart, and a sample of diaphyseal bone marrow 
from one of the femurs contained approximately 54% 
fat, as measured by comparison of the wet and dry 
weights of the sample. A large area of the skin along 
the ventral side of the neck, about 50 cm?, was miss- 
ing and had been replaced by scar tissue that released 
a foul odour indicative of suppuration. The portion of 
the cable along the ventral side of the neck had tran- 
sected the full diameter of the trachea and was 
embedded in scar tissue between the trachea and the 
esophagus (Figure 1). The wall of the severed portion 
of the trachea was completely healed, although the 
tissue of repair had extended as a thin circular band 


243 


244 


into the tracheal lumen, reducing its surface area to 
about one-seventh of its original size (Figure 2). The 
esophageal lumen was patent. There was no gross evi- 
dence of suppuration accompanying the scar tissue that 
surrounded the trachea and the cable. At the time of 
necropsy, the cable was freely movable within the 
affected tissues. The architecture of both jugular veins 
disappeared at the level of the scar tissue. A thin probe 
inserted into the cranial portion of the common caro- 
tid artery on both sides of the trachea was blocked 
from further progression at this level. The outline of 
the vagosympathetic trunk on both sides was also lost 
among the scar tissue. No other significant lesion was 
seen at necropsy. Samples of the affected portion of 
trachea and surrounding soft tissues were fixed in for- 
malin, processed routinely for microscopic examina- 
tion, and stained with hematoxylin and eosin. 

Microscopically, the scar tissue surrounding the 
affected portion of trachea consisted of abundant dense 
fibrous tissue, much of which was birefringent under 
polarized light. This fibrous tissue enclosed several 
pockets of necrotic material (including debris of in- 
flammatory cells) and, in some areas, contained numer- 
ous cross sections of hair shafts. Serial sections of this 
tissue, parallel to the length of the trachea, revealed 
on both sides an artery, about 3 mm in diameter, and 
a nerve, about 1 mm in diameter. Taking into account 
the shrinkage that normally occurs in tissues processed 
for microscopic examination, the diameters of these 
two structures were compatible with those of the com- 
mon carotid artery and vagosympathetic trunk, respec- 
tively, for an animal of this size. Depending on the 
section examined, the lumen of the common carotid 
artery on both sides of the trachea was either empty 
or partly or completely filled with a well organized 
thrombus. The vagosympathetic trunk on both sides 
was morphologically normal, except for a few degen- 
erative lesions suggesting some loss of axons and their 
myelin sheaths (wallerian degeneration). The tissue 
within the tracheal lumen also consisted of dense fi- 
brous tissue that enclosed some necrotic material and 
numerous cross sections of hair shafts. 


Discussion 

Ideally, a snare should cause rapid death by neck 
strangulation and subsequent asphyxiation. A quick 
death is more humane and also gives the animal less 
time to find a way out of the snare. A variety of factors 
can influence the performance of a snare. Some lock- 
ing mechanisms tighten less quickly and firmly than 
others (Phillips 1996), or small kinks in the cable may 
weaken it and make it easier to break or may prevent 
full function of the lock. Inclusion of a swivel attach- 
ing the lock to the cable (not present in this case) can 
prevent the cable from kinking as the lock slides 
along it. If the cable is not anchored firmly enough, 
the animal may be able to pull off the snare before 
maximum closure has been achieved. If death is not 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


rapid and the cable is anchored too low, it may be more 
easily accessible to the animal’s jaws and be chewed 
through. Seventeen of 131 (13%) Coyotes caught by 
one of three types of snares studied by Phillips (1996) 
managed to escape, one by breaking the lock and 16 by 
chewing through the cable. 

In this Coyote, the relative abundance and density 
of the scar tissue in the neck and, microscopically, its 
birefringence under polarized light suggested that the 
failed capture had occurred at least a few weeks prior 
to death. The trapping season for Coyote and Red Fox 
(Vulpes vulpes) on PEI officially terminated on Janu- 
ary 15, more than a month before this animal was 
found. The reason why it managed to escape capture 
was not determined, although the portion of cable dan- 
gling from its neck was of the right length to suggest 
that the animal had chewed through it rather than pull 
the whole cable from where it was anchored. Unfortu- 
nately, the animal’s gums and teeth were not examined 
for the presence of recent damage that would have sup- 
ported this suggestion. At the time of escape, sufficient 
closure of the snare may have already been achieved 
for the cable to exert a strong pressure on soft tissues 
of the ventral side of the neck and be forced gradually 
through these tissues over the following weeks. Grad- 
ual transection and concurrent repair of the tracheal 
wall would thus have allowed it to remain continuous. 
Alternatively, transection of all soft tissues, including 
the trachea, could have occurred at the time of the 
animal’s initial struggle in the snare. Such rapid tran- 
section of the trachea, however, would more likely 
have resulted in the formation of a gap between its 
cut ends, thus preventing them from healing together. 
Moreover, if the trachea had been rapidly transected, 
the jugular veins would probably also have been cut 
and the animal would presumably have bled to death. 
The thicker hair coat and skin on the dorsal side of the 
neck as compared to the ventral side (Scott et al. 2001) 
and the tough nuchal ligament on the dorsal side (Evans 
1993) may have prevented the cable from moving into 
the tissues from that side. 

The common carotid arteries and the vagus nerves 
are important structures running along both sides of 
the trachea. In this animal, the lumen of both common 
carotid arteries was occluded. However, this is not a 
critical problem, at least in canids. In dogs, both com- 
mon carotid arteries can be ligated experimentally 
without causing any clinical signs, implying that col- 
lateral circulation (e.g., via the vertebral arteries) is 
able to quickly compensate and transport sufficient 
blood to the brain (Whisnant et al. 1956). Such col- 
lateral circulation would have even more time to devel- 
op if the obstruction were gradual, as is suspected to 
have occurred in this case. The same reasoning applies 
to the jugular veins, located much more superficially 
in the ventral region of the neck, since collateral cir- 
culation within the venous system can develop even 
more readily than in the arterial side. Proper function 


2004 


DAOUST AND NICHOLSON: NECK INJURY IN A COYOTE 245 


FiGure |. Right lateral view of the neck of a Coyote. The carcass is lying on its back, with the head to the left. The cable 
from a snare has cut through the trachea (T) and is now lodged between it and the esophagus (E). The distortion of 
the wall of the trachea just above the cable as been caused by scar tissue of repair. A metal rod has been inserted in 
the lumen of the esophagus. The common carotid artery (C) and vagosympathetic trunk (V) are clearly visible to the 
left of the cable. The lock of the snare is at the bottom of the picture. (Bar = 2 cm.) : 


of at least one of the vagosympathetic trunks is, how- 
ever, essential to life. The vagal (parasympathetic) 
portion of these nerves originates in the brain and 
supplies efferent and afferent innervation to many 
vital organs, including the cardiovascular system and 
digestive tract (Andrews and Lawes 1992; Esler 1992; 
Guyton and Hall 1996). In particular, vagal stimula- 
tion protects against excessive stimulation of cardiac 
function by the sympathetic nervous system during 
physical activity and stress. In this Coyote, it was not 
possible to dissect the vagosympathetic trunk on either 
side of the affected portion of trachea, because both 
trunks were well embedded in scar tissue at this level. 
However, careful microscopic examination showed 
that both trunks were in large part morphologically in- 
tact. Although their structure had likely been stretched, 
and their path distorded, by the cable, the gradual pro- 
cess through which this happened would presumably 
have provided sufficient time for anatomic and physio- 
logical compensation. 

The exact cause of the eventual demise of this coy- 
ote was not clear. There are undocumented reports by 
veterinary practitioners of collars being left permanent- 
ly on growing dogs and eventually becoming buried 
under the skin, with damage limited to chronic sup- 
puration of the subcutis (J. B. Miller, Department of 
Companion Animals, Atlantic Veterinary College, Uni- 
versity of PEI, personal communication). Similarly, a 


square of a monofilament gillnet mesh was found al- 
most totally embedded in the snout of an otherwise 
healthy Porbeagle (Lamna nasus) that had been caught 
on the Scotian Shelf (Benz et al. 2001). However, the 
marked reduction in size of the tracheal lumen in this 
Coyote may have reduced its stamina and prevented it 
from foraging efficiently. The esophageal lumen was 


FiGurRE 2. Cranial view of the trachea of a Coyote at the level 
where the cable of a snare has cut through this organ. 
The ventral side of the trachea is at the top. The carti- 
laginous ring (CR) outlines what should be the normal 
size of the tracheal lumen. Scar tissue of repair has 
greatly narrowed this lumen. (Bar = | cm.) 


246 


patent, but the presence of the cable against its ventral 
surface may have interfered with the passage of large 
food boluses. Albeit seemingly superficial, the suppu- 
rative process and, therefore, probable bacterial in- 
fection, associated with the soft tissue damage would 
have further drained the animal’s energy reserves. 
Although this animal had not yet reached a stage of 
emaciation, its poor body condition, combined with 
the energy demands imposed by the winter season, 
may have been sufficient to bring it to a stage of 
exhaustion, perhaps combined with hypothermia. The 
clinical significance, if any, of the mild damage seen 
microscopically in the vagosympathetic trunks could 
have been assessed only in the live animal. 

The efficacy of various trapping devices, including 
snares, can now be properly tested under standardized 
laboratory conditions (Proulx and Barrett 1990, 1991). 
Ultimately, however, the proportion of animals that 
fail to die rapidly and undergo prolonged suffering 
depends much on the experience and expertise of the 
trappers in the field. Regardless of the exact cause of 
failure of the snare in this case, its outcome emphasizes 
the need for continued attention to appropriate train- 
ing of hunters and trappers, in order to ensure that the 
best trapping devices and capture techniques are used 
consistently and, as a result, as few animals as pos- 
sible undergo undue suffering. It also illustrates the 
extreme resilience of Coyotes. 


Acknowledgments 
This article benefited greatly from discussions with 
Neal Jotham. 


Documents Cited [marked * in text] 

Agreement on International Humane Trapping Standards. 
The European Community, the Government of Canada, 
and the Government of the Russian Federation. 1997. 
Department of Foreign Affairs and International Trade, 
Ottawa, Ontario, Canada. 31 pages. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


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Neuroanatomy and physiology of abdominal vagal afferents. 
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Press, Boca Raton, Florida, 316 pages. 

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Boddicker, M. L. 1982. Snares for predator control. Pro- 
ceedings of the Vertebrate Pest Conference 10: 50-54. 
Esler, M. 1992. The autonomic nervous system and cardiac 
arrhythmias. Clinical Autonomic Research 2: 133-135. 
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anatomy of the dog. Edited by H. E. Evans. W. B. Saunders 

Company, Toronto, 3" edition, 1113 pages. 

Guthery, F. S., and S. L. Beasom. 1978. Effectiveness and 
selectivity of neck snares in predator control. Journal of 
Wildlife Management 42: 457-459. 

Guyton, A. C., and J. E. Hall 1996. Textbook of medical 
physiology. W. B. Saunders Company, Toronto, 9" edition: 
pages 107-119, 793-802. 

Phillips, R. L. 1996. Evaluation of 3 types of snares for cap- 
turing coyotes. Wildlife Society Bulletin 24: 107-110. 
Proulx, G., and M. W. Barrett. 1990. Assessment of power 
snares to effectively kill red fox. Wildlife Society Bul- 

letin 18: 27-30. 

Proulx, G., and M. W. Barrett. 1991. Ideological conflict 
between animal rightists and wildlife professionals over 
trapping wild furbearers. Transactions of the North Ameri- 
can Wildlife and Natural Resources Conference 56: 
387-399. 

Scott, D. W., W. H. Miller Jr., and C. E. Griffin. 2001. 
Muller & Kirk’s small animal dermatology. W. B. Saunders 
Company, Toronto, 6" edition, pages 1-70. 

Whisnant, J. P., C. H. Millikan, K. G. Wakim, and G. P. 
Sayre. 1956. Collateral circulation to the brain of the dog 
following bilateral ligation of the carotid and vertebral 
arteries. American Journal of Physiology 186: 275-277. 


Received 7 February 2003 
Accepted 26 October 2004 


Interactions of Brown Bears, Ursus arctos, and Gray Wolves, Canis 
lupus, at Katmai National Park and Preserve, Alaska 


Tom S. SMITH*, STEVEN T. PARTRIDGE”, and JOHN W. SCHOEN? 


4 United States Geological Survey Alaska Science Center, 1011 E. Tudor Road, Anchorage, Alaska 99503 USA 
>Alaska Office — National Audubon Society, 308 G. Street, #217, Anchorage, Alaska 99501 USA 


Smith, Tom S., Steven T. Partridge, and John W. Schoen. 2003. Interactions of Brown Bears, Ursus arctos, and Gray Wolves, 
Canis lupus, at Katmai National Park and Preserve, Alaska. Canadian Field-Naturalist 118(2): 247-250. 


We describe several encounters between Brown Bears (Ursus arctos) and Gray Wolves (Canis lupus) that were observed at 
Katmai National Park and Preserve in southwest Alaska. Katmai Brown Bears and Gray Wolves were observed interacting 
in a variety of behavioral modes that ranged from agonistic to tolerant. These observations provide additional insight regard- 
ing the behavioral plasticity associated with bear-wolf interactions. 


Key Words: Brown Bear, Ursus arctos, Gray Wolf, Canis lupus, interactions, Katmai National Park and Preserve, Alaska. 


Flexibility in mammalian behavior parallels the evo- 
lution of large brains (Gilbert 1989). Gittleman (1986) 
presented data showing that bears and wolves have 
the highest brain:body weight ratios within the order 
Carnivora, and claimed that this accounts for their 
behavioral plasticity. The context of the encounter set- 
ting, as well as the behavior of individual animals, like- 
ly influences the resulting relationship between bears 
and wolves at any given time. 

Alaska Brown [Grizzly] Bears (Ursus arctos) and 
Gray Wolves (Canis lupus) are sympatric over much 
of their range and often compete for access to the 
same food resources (Adams et al. 1995). As sym- 
patric, apical predators, bears and wolves probably 
interact frequently, although reports in the scientific 
literature are scant. The accounts reported illustrate 
the variable nature of Brown Bear — Gray Wolf inter- 
actions, and range from mutual tolerance (Lent 1964), 
to competitive (Adams et al. 1995; Hornbeck and 
Horejsi 1986; Ballard 1982), to predatory (Ballard 
1980; Hayes and Mossop 1987). This note describes 
additional Brown Bear — Gray Wolf interactions ob- 
served at Katmai National Park and Preserve on the 
Alaska Peninsula, approximately 400 km southwest 
of Anchorage, Alaska. 


Wolves harass bears 

On 24 June 2001, while conducting Brown Bear 
research at Hallo Bay (154° 05’ W, 58° 27’ N) on the 
Katmai coast, the authors observed a pack of five 
wolves harass a pair of courting Brown Bears on the 
edge of extensive salt marsh meadows (Carex spp., 
Plantago spp., Triglochin spp.). Prior to this observa- 
tion, we had repeatedly heard wolves “chorus howI- 
ing” (Harrington and Asa 2003), and noted that the 
pack was gradually nearing our research field camp 
situated on the far end of the meadow. Anticipating 
their arrival, we positioned ourselves where we could 
observe them when they entered the meadow. Although 


it was late in the evening (2230 h Alaska Daylight 
Time), it was still light enough to see so we patiently 
waited while scanning the meadow. About 400 m south 
of our position, a pair of Brown Bears foraged on 
sedges in the meadow near a willow-alder (Salix spp., 
Alnus spp.) thicket. After we waited 10 minutes, the 
wolf pack suddenly appeared, emerging from the dense 
thicket about 25 m north of the two bears. The wolves 
trotted single file in the direction of the bears. Three 
of the five wolves had black-gray coats, similar in pat- 
tern to that of the German Shepherd (C. domesticus) 
breed of domestic dog; one wolf’s coat was pure white; 
the other coal black. All five wolves appeared to be 
mature adults, standing nearly 1 m at the shoulder and 
similar in stature. 

As one, the wolves circled south from the meadow’s 
center toward the bears, which by then were focused 
on the wolves. The five wolves lined up, head-to-head, 
with the large male Brown Bear with < 2 m separating 
the lead wolf and the bear. Suddenly, one wolf rushed 
the maie bear, which in turn lunged and swatted at the 
attacking wolf with its foreleg. Leaping forward, duck- 
ing the bear’s defensive swat, then breaking away, the 
wolves took turns harassing the male bear. The bear 
responded to the attack with short lunges and aggres- 
sive paw swats while slowly backing away. About a 
minute into the confrontation, two wolves circled the 
bear and lunge-nipped at his unprotected hindquarters. 
In defense, the male bear swiftly spun around and swat- 
ted at the closest attacker, but the wolf dodged the bear’s 
swinging paw then trotted a few meters away, tail wag- 
ging high. While the male bear counter-attacked the 
wolves formerly at his rear, the other three wolves, now 
at his unprotected hindquarters also began a nipping 
attack, to which the bear again whirled around, lunged, 
growled, and aggressively swatted at the wolves. The 
wolves easily outmaneuvered the large bear’s paw 
swipes and lunges with quick, evasive maneuvers. 


The wolves and male bear repeated this attack-count- 
er attack interaction several times before a group of 
three wolves split from the pack and rushed the female 
bear that had been watching from a distance of about 
10 meters, near the edge of the thicket. Repeating the 
same lunge, nip, and run strategy, the two groups of 
wolves attacked each bear, which rapidly spun around 
to defend themselves from the lunging-biting wolves. 
At this point, approximately 10-20 meters separated 
the two bears as they fended off the wolves. During 
these aggressive interactions, neither wolves nor bears 
were observed making physical contact, although the 
confrontation appeared quite serious. After 5 minutes 
of sustained confrontation, two wolves abruptly termi- 
nated their attack and began chasing each other in tight 
circles a short distance from the other wolves, wag- 
ging tails held high. Moments later, the remaining three 
wolves also withdrew, excitedly darted about, wagging 
tails, and then joined the others. After regrouping about 
30 m from the bears, all wolves participated in a series 
of long, drawn-out “chorus” howls. From the forest 
to the west, responding howls (possibly from a single 
wolf) were heard in apparent reply. Moments later, 
the pack trotted southward into the rapidly dimming 
twilight, away from the bears and researchers. The 
entire interaction took approximately 20 minutes from 
the time the wolves were observed entering the mead- 
ow to when they disappeared into the dusk. 

To the best of our knowledge, there had been no 
food source from which the wolves were attempting 
to drive the bears. It appeared to us that this wolf pack 
had encountered and approached these two bears and 
attempted to test, or harass, them for a short period 
before breaking off the encounter and moving on. 


Wolves steal fish from bears 

In mid-July 2001, a pair of Gray Wolves was ob- 
served stealing Chum Salmon (Oncorhynchus keta) 
from Brown Bears at Middle Creek where it empties 
into Hallo Bay on the Katmai coast. National Park 
Service (NPS) ranger Stephens Harper first saw the 
wolves when they trotted past the rangers’ camp at 
0830 h, heading to the nearby beach. Bears and wolves 
frequent the tidal flats because the stream channel is 
quite shallow at low tide, thus making salmon more 
vulnerable to predation. Harper walked to the stream 
mouth a few minutes before noon and observed two 
wolves and five single adult bears fishing in close 
proximity to one another. 

Both wolves were gray with black highlights; one 
appeared mature, while the other was younger, lankier, 
and smaller (Figure 1). Fishing at the river mouth were 
five adult single bears. The younger wolf bedded on a 
gravel bar some distance from the bears while the other, 
much closer, visually scanned them, apparently wait- 
ing for one to catch a fish successfully. Once a bear 
caught a fish, the older wolf would move in, head low, 
and attempt to steal it by suddenly lunging and snag- 
ging a portion away from the bear. In response, the 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


bear would protect its quarry by bolting and running 
after the wolf. Although there was a lot of chasing up 
and down the stream banks, the wolf easily outran and 
outmaneuvered these bears. This strategy worked, 
however, in getting bears to abandon what was left of 
the fish and the wolf would circle back around and eat 
these scraps. In turn, the older wolf would trot over 
to where the other was bedded, regurgitate a portion, 
and then head back to where the bears were fishing. 
During the entire hour that Harper watched, the wolf 
determinedly attempted to snatch fish from bears. 
Although these wolves attempted to catch salmon for 
themselves on several occasions, they were unsuccess- 
ful. One wolf, however, scavenged a Starry Flounder 
(Platichthys stellatus) lying on the beach that had 
been exposed at low tide. 


Wolf displaces a bear from a moose carcass 
In June 1997, an NPS employee observed a single 
wolf feeding on a Moose (Alces alces) carcass near a 
gravel pit adjacent to the Valley of Ten Thousand 
Smokes Road, approximately 13 km from Brooks 
Camp. Adjacent to the gravel pit is a wetland-pond 
complex impounded by a Beaver (Castor canadensis) 
dam. The following day, NPS employees Jim Gavin and 
Tom Ferguson observed a wolf and bear simultaneous- 
ly at the site. The wolf was gray with black highlights, 
appeared mature, but thin. The bear was a young fe- 
male, approximately 4 years of age. Initially, only the 
bear fed on the carcass while the bedded wolf watched 
from 40 m distant. After a while, the wolf moved in 
close and began harassing the bear by charging and 
biting at its hind legs. The wolf’s constant attacks dis- 
rupted the bear’s feeding as it defended itself. After 
1.5 min of intense harassment, the bear took flight into 
the forest and the wolf usurped the carcass and started 
feeding. After approximately 10 min of feeding, the 
satiated wolf trotted off into the forest, reappeared 
20 min later, then repeated the process several times, 
likely caching food (L. Adams, USGS research biol- 
ogist, personal communication), or transporting it to 
young at a den nearby. Gavin observed the wolf feed 
in this manner for three consecutive days. On the third 
day, while the wolf was absent, the bear returned and 
resumed control of the Moose carcass. The wolf re- 
appeared but did not approach any closer than 40 m 
to the feeding bear. Gavin did not observe the wolf 
attempt to displace the bear as before, although his 
observation sessions were only 30 min daily. 


Bears and wolves fishing together 

Chum Salmon enter freshwater to spawn in Hallo 
Bay’s Middle Creek from early July through August. 
In 2001, Brown Bears and wolves were repeatedly 
observed fishing side-by-side (approximately 5 m apart) 
at this location with no apparent interactions or obvious 
concern toward the other species. Like bears, wolves 
sat patiently on the gravel bank and scanned the water 
for incoming salmon. Upon catching sight of an incom- 


2004 


ing salmon, both bears and wolves would leap into the 
stream and attempt to capture it. Both bears and wolves 
appeared indifferent to one another, completely fo- 
cused on fishing. 

Similarly, in August 1992, NPS rangers B. Holmes 
and S. Klenzendorf observed a lone wolf and several 
bears fishing in close association for 2 h at Margo 
Creek (155° 03’ W, 58° 29’ N) in the western portion 
of Katmai National Park. At 1210 h on 13 August, an 
old female Brown Bear was fishing when a wolf ap- 
peared suddenly on the riverbank, watched the bear, 
and then sat down. The wolf was moderate in size and 
of gray coloration. The wolf wandered in the direction 
of the bear, entered the creek and retrieved a spawn- 
ing Red Salmon (Oncorhynchus nerka). As the wolf 
fed, the bear wandered downstream towards the wolf. 
Consequently, the wolf abandoned its catch, moved 
quickly away, picked up another salmon, and then 
moved into the nearby White Spruce (Picea glauca) 
forest. At 1245 h, the wolf reappeared from the forest, 
plunged into the creek and emerged with a live salmon. 
Carrying the salmon in its mouth, the wolf disappeared 
again into the heavy cover of the forest. At 1255 h, the 
wolf reappeared and entered the creek in search of sal- 
mon. As the wolf waded about searching for fish, the 
same bear waded downstream until she and the wolf 
were about 20 m apart. In apparent response, the wolf 
abruptly ceased fishing, climbed out of the river and 
sat and watched the bear as it sought fish in that section 
of stream. Twenty minutes later, the wolf moved up- 
stream and caught a salmon. As the wolf fed, a large 
adult male bear suddenly appeared thus causing the 
wolf to leave the area. The large male bear quickly 
caught a fish, then left the river, at which time the wolf 
reappeared. Moments later, the wolf caught a fish then 
reentered the forest. The observers left at 1402 h, im- 
pressed with the degree of tolerance the two species 
appeared to have for one another. 


Wolf and bears traveling together 

In June 1989, Aleska Szweda, an NPS employee 
stationed at Brooks Camp (155° 05’ W, 58°33 ‘ N) in 
the western portion of Katmai National Park, observed 
a lone wolf traveling in the company of two Brown 
Bears. Szweda was driving along the park road, approx- 
imately 8 km from Brooks Camp, when she noticed 
what appeared to be two bears in the middle of the 
roadway, approximately 400 m ahead. The lumbering 
vehicle’s approach prompted the bears to abandon the 
roadway suddenly, exposing a wolf that had appar- 
ently been in their immediate company (<2 m). The 
wolf too left the road, following the bears into the 
forest. 

The following day, five park visitors reported see- 
ing two bears and a wolf cross the Brooks River at a 
location known locally as the Oxbow. It was generally 
believed that these were the same three observed by 
Szweda the day previous along the park road, 8 km 
distant. The bears and wolf were clearly traveling as 


SMITH, PARTRIDGE, and SCHOEN: INTERACTIONS OF BEARS AND WOLVES 


249 


FIGURE 1. Gray Wolf at Hallo Bay, Katmai National Park and 
Preserve, Alaska (Kent Fredriksson photograph). 


a group as they worked their way down the riverbank, 
probed the riverbed for the shallowest crossing, and 
then emerged on the other side. It was reported that 
the trio fed together in an unusual manner; when the 
bears caught fish they fed only upon the brains and 
skin, leaving the rest for the wolf which, sitting im- 
mediately adjacent, would spring forward, snatch and 
eat the remainder. It is not unusual to see Red Foxes 
(Vulpes vulpes) or gulls (Larus spp.) attempting to 
snatch fish scraps from Brown Bears on salmon streams. 
But bears in these instances are far from cooperative, 
often lunging at the would-be thief, catching and killing 
them on occasion (T. Smith, personal observation). 
However, in this case, the bears tolerated the wolf’s 
presence and, according to observers, did nothing to 
discourage its very close (1 m) proximity to them. 


Acknowledgments 

We gratefully acknowledge the contributions of 
Brian Holmes, Ronald Squibb, Tamara Olson, Kerry 
Sullivan, Jim Gavin, Aleska Szweda, Stephens Harper, 
and Layne Adams. We also thank Rick Clark and Troy 


Hamon of Katmai National Park for funding Brown 
Bear research in the park. 


Literature Cited 

Adams, L. G., F. J. Singer, and B. W. Dale. 1995. Caribou 
calf mortality in Denali National Park, Alaska. Journal of 
Wildlife Management 59: 584-594. 

Ballard, W. B. 1980. Brown Bear kills Gray Wolf. Canadian 
Field-Naturalist 94: 91. 

Ballard, W. B. 1982. Gray wolf-brown bear relationships in 
the Nelchina Basin of south-central Alaska. Pages 71-80 
in Wolves of the world: perspectives of behavior, ecology, 
and conservation. Edited by F. H. Harrington and P. C. 
Paquet. Noyes Publications, Park Ridge, New Jersey. 

Gilbert, B. K. 1989. Behavioral plasticity and bear-human 
conflicts. Pages 1-8 in Bear-People Conflicts: Proceedings 
of a Symposium on Management Strategies. Edited by 
M. Bromley. Yellowknife, Northwest Territories. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Gittleman, J. L. 1986. Carnivore brain size, behavioral ecol- 
ogy and phylogeny. Journal of Mammalogy 67: 23-36. 
Harrington, F. H., and C. S. Asa. 2003. Wolf communica- 
tion. Pages 66-103 in Wolves: behavior, ecology and con- 
servation. Edited by L. David Mech and Luigi Boitani. 

The University of Chicago Press. Chicago, Illinois. 

Hayes, R. D., and D. H. Mossop. 1987. Interactions of wolves, 
Canis lupus, and brown bears, Ursus arctos, at a wolf den 
in the northern Yukon. Canadian Field-Naturalist 101: 603- 
604. 

Hornbeck, G. F., and B. L. Horejsi. 1986. Grizzly bear, 
Ursus arctos, usurps wolf, Canis lupus, kill. Canadian Field- 
Naturalist 100: 259-260. 

Lent, P. C. 1964. Tolerance between grizzlies and wolves. 
Journal of Mammalogy 45: 304-305. 


Received 17 February 2003 
Accepted 2 December 2003 


Wild Turkey, Meleagris gallopavo silvestris, Behavior in Central 
Ontario During Winter 


Linu P. Ncuyen,! JOSEF HAMR,” and GLENN H. PARKER! 


'Department of Biology, Laurentian University, Ramsey Lake Road, Sudbury, Ontario P3E 2C6 Canada 

*Northern Environmental Heritage Institute, Cambrian College of Applied Arts and Technology, 1400 Barrydowne Road, 
Sudbury, Ontario P3A 3V8 Canada 

3Present address: Watershed Ecosystem Graduate Program, Trent University, 1600 West Bank Drive, Peterborough, Ontario 
K9J 7B8 Canada 


Nguyen, Linh P., Josef Hamr, and Glenn H. Parker. 2004. Wild Turkey, Meleagris gallopavo silvestris, Behavior in Central 
Ontario During Winter. Canadian Field-Naturalist 118(2): 251-255. 


Home range size, food habits, and roost site selection are described for the Eastern Wild Turkey (Meleagris gallopavo 
silvestris) introduced on the Precambrian Shield in central Ontario during the winters 1999 and 2000. Monthly home range 
size was correlated primarily to snow depth, although it was also likely associated to other factors, including food avail- 
ability and/or roost site availability. Ferns and allies were used more than available, whereas monocots were used less than 
expected. Roost site-selection was primarily influenced by tree height. If the Eastern Wild Turkey is to expand its northern 


range in Ontario, winter food and roost site availability may be the primary determinants for successful introductions. 


Key Words: Eastern Wild Turkey, Meleagris gallopavo silvestris, home range size, roost site, food availability, Ontario. 


The Eastern Wild Turkey (Meleagris gallopavo sil- 
vestris) is a large, highly adaptable gallinaceous bird 
that is common throughout southern Ontario as a result 
of introductions in the last 20 years (Bellamy 2001). 
For northern turkey populations, winter is the most 
stressful season for satisfying energy demands (Ober- 
lag et al. 1990). Variations in home range size and re- 
source selection appear to be governed by snow depth 
(Porter et al. 1980; Vander Haegen et al. 1989) and, 
to a lesser extent, ambient temperature (Oberlag et al. 
1990). Numerous accounts of turkey winter biology 
(e.g., Glover and Bailey 1949; Wunz and Hayden 1975; 
Porter et al. 1980; Kilpatrick et al. 1988; Vander Haegen 
et al. 1989), refer primarily to studies conducted in the 
United States. The winter biology of turkeys in these 
southern habitats may be different from that of birds 
living in Ontario due to inherent differences in climate 
and habitat. Hence, we report on home range size, 
food habits, and roost site use by Eastern Wild Turkeys 
introduced to the Precambrian Shield in central Ontar- 
io during the winters 1999 and 2000. It was hypothe- 
sized that turkey movement and forage availability were 
inversely related to snow depth. It was also hypothe- 
sized that the tallest trees provided optimum wild tur- 
key roost sites due to their sturdiness. 


Study Area 

This study was conducted from November to March 
1999/2000 and 2000/2001 near Noélville, approximate- 
ly 60 km southeast of Sudbury, Ontario (46°10'N, 
80°25'W). The 169-km/? study area was located within 
the Great Lakes—St. Lawrence Ecotonal Forest Region 
(Rowe 1972), characterized by flat to rolling topog- 
raphy, interrupted by rock outcrops and narrow valleys. 
The habitat consisted of 20% softwood forests, 37% 


hardwood forests, 28% abandoned pasture and hay- 
fields, and 15% residential areas and rock outcrops. 
Beef farming was the dominant land use, with many 
fields cultivated for corn silage or pasture grasses. 
The forested areas were dominated by White Birch 
(Betula papyrifera) and Trembling Aspen (Populus 
tremuloides), interspersed with Balsam Fir (Abies 
balsamea), Eastern White and Red Pine (Pinus .strobus, 
P. resinosa), Red Oak (Quercus rubra), White Spruce 
(Picea glauca), Red and Sugar Maple (Acer rubrum, 
A. saccharum), and Eastern Hemlock (Tsuga canaden- 
sis). Shrubs included raspberries (Rubus spp.), Bracken 
Fern (Pteridium aquilinum), blueberries (Vaccinium 
spp.), Beaked Hazel (Corylus cornuta), and asters 
(Aster spp.). 

January, the coldest month of the year, had a mean 
temperature of —13.9°C and —10.5°C in 2000 and 2001, 
respectively. Total snowfall was 216.0 cm in 1999- 
2000 (10.0% below the 30-year norm) and 328.3 cm 
in 2000-2001 (22.2% above the 30-year norm). Snow 
depths exceeded 25 cm for 38 days in 1999-2000 and 
111 days in 2000-2001. 


Methods 
Capture and Radio-Tracking 

Wild Turkeys were captured with rocket nets (Hawk- 
ins et al. 1968) in southern Ontario and upper New 
York for introduction to the study area in February 
and March 1999 (10 males and 26 females) and 
March 2000 (13 females). Female turkeys were fitted 
with backpack-style, 32.5 g (1% of mean body mass), 
mortality-mode VHF radio-transmitters (Holohil Sys- 
tems Ltd., Carp, Ontario). We tracked radio-fitted birds 
two to four days per week, or until battery failure, from 
1999 to 2001. Locations of birds were determined by 


251 


Diy) 


triangulation using >3 locations (Heezen and Tester 
1967) taken less than 15 minutes apart with a 2-element 
H antenna and portable receiver-scanner (Model STR- 
1000, Lotek Engineering Inc., Newmarket, Ontario). 
Average telemetry error was 156.9 m + 21.1 SE (n= 
40). 


Home Range 

Monthly home range sizes were calculated using 
the 100% minimum convex polygon method (White 
and Garrott 1990). All spatial analyses were conducted 
using an ArcView GIS software (Environmental Sys- 
tems Research Institute, Redlands, California), with the 
Animal Movement Analysis (Hooge and Eichenlaub 
1997) and Spatial Analyst Extensions. 


Forage Selection 

Active turkey feeding areas were identified by tracks, 
and forage plants were identified and collected. Crop 
contents from dead specimens provided a supplemen- 
tal source of information. Forage species eaten were 
recorded regardless of the number of bites using a 
modified point-quarter method (Jost et al. 1999) during 
2000-2001. Point samples of available plants adjacent 
to turkey tracks in the herb (less than 0.5 m) and shrub 
(0.5 to 2.0 m) layers were selected randomly and 
identified. Plants were grouped into five classes: (1) 
mosses, (2) ferns and allies, (3) conifers and allies, 
(4) monocots, and (5) dicots. 


Roost Site 

Roost sites were found by (1) observing commonly 
used trees, (2) finding turkey droppings under trees 
(Hoffman 1968), (3) examining areas where Wild Tur- 
keys were common after snowfalls, or (4) locating 
radio-fitted birds on trees before sunset. Locations of 
roost sites were determined using a Global Positioning 
System (GARMIN International Inc., Olathe, Kansas). 
Equal numbers of random trees that may have pro- 
vided potential roost sites within our study area were 
generated in ArcView GIS. Random trees were located 
using a Global Positioning System, and trees that had 
diameter at breast height (dbh) less than 10.2 cm were 
discarded (Kilpatrick et al. 1988). 

Elevation, tree dbh, percent canopy cover, canopy 
density, distance to habitat edge, distance to open water, 
and tree height for actual and selected trees were mea- 
sured. Elevation was measured with an altimeter, tree 
dbh with a diameter tape, percent canopy cover by 
averaging readings at each compass directions (N, S, 
W, E) using a convex spherical densiometer, canopy 
density by counting tree trunks of dbh =10.2 cm with- 
in 5 m of actual roost sites, and tree height from esti- 
mates on a subjectively selected “average” tree. Dis- 
tances to habitat edge and open water were also deter- 
mined by plotting roost sites on Ontario Base Maps 
(OBM) and Forest Resource Inventory (FRI) maps. 
The used habitat variables either had been previously 
described in the literature (Tzilkowski 1971; Kilpatrick 
et al. 1988; Chamberlain et al. 2000), or were hypoth- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


esized correlates based on the winter biology of the 
species. 


Statistical Analyses 

Data were pooled to maintain sufficient sample 
size, unless otherwise noted (Alldredge and Ratti 1986; 
Thomas and Taylor 1990). Home range sizes were 
compared between years using the Mann-Whitney U- 
test. Monthly home ranges were compared using the 
Kruskal Wallis test (Zar 1999). Two measurements 
used to examine the effects of winter on monthly home 
range size were (1) number of days with snow depths 
over 25 cm (Porter et al. 1980), and (2) number of 
days with minimum temperature less than —16.2°C 
(Oberlag et al. 1990). Spearman rank-order correlation 
analyses were used to compare mean monthly home 
range size with these two variables. The number of 
days with snow depth over 25 cm and the number of 
days with minimum temperature less than —16.2°C 
were also compared between years using the Mann- 
Whitney U-test. 

Chi-square analysis was used to compare the fre- 
quencies of plants browsed by turkeys to availability 
(Neu et al. 1974). When forage selection differed, a 
Bonferroni Z-test was used to identify plants that 
were browsed more or less than availability (Byers et 
al. 1984). A series of one-way analyses of variance 
(ANOVA) was used to compare each habitat variable 
associated with roosting and random sites. Pearson cor- 
relation analyses were performed for the significant 
habitat variables. In order to minimize artifacts from 
environmental variations in habitat characteristics, an 
analysis of covariance (ANCOVA) was used to deter- 
mine significant habitat variables. All analyses were 
conducted using Statistical Package for the Social 
Sciences (SPSS Inc., Chicago, Illinois) with signifi- 
cance level set at a = 0.10. 


Results 
Home Range 

We obtained 441 locations from 12 radio-fitted 
turkeys from November 1999 to March 2000 and 
from November 2000 to January 2001. Mean winter 
home range size differed between years (U = 142.0, 
P =0.002), as did monthly home range size (Novem- 
ber: U = 9.0, P = 0.083; December: U = 8.0, P = 0.060: 
January: U = 3.0, P = 0.020). In addition, home range 
size differed among months in the two years (c? = 7.79, 
df = 4, P = 0.100; Table 1). Monthly home range size 
was correlated with the number of days with snow 
depth more than 25 cm (7, = —0.71, n= 59, P = 0.050) 
and the number of days with minimum temperature 
less than —16.2°C (r, = 0.75, n= 59, P = 0.031). 

The number of days with snow depth over 25 cm 
differed (U = 4.0, P = 0.095) between 1999-2000 
(9.60 + 5.91 days, n = 5) and 2000-2001 (22.6 + 4.88 
days, n = 5), while the number of days with mini- 
mum temperature less than —16.2°C did not differ 
(U = 10.5, P = 0.690) between 1999-2000 (10.00 + 


2004 NGUYEN, HAmr, and PARKER: WILD TURKEY BEHAVIOR 255 


TABLE |. Winter home range size (ha) of Eastern Wild Turkey hens in Noélville, near Sudbury, Ontario, 1999-2001 (SE = 
standard error of the mean). 


1999-2000 2000-2001 1999-2001 (Pooled Data) 
n Mean SE n Mean SE n Mean SE 

Winter 9 249 60 4 58 19 13 204 47 
Monthly 
November 9 399 198 5 90, 54 14 288 132 
December 9 145 60 5) 4] 8 14 108 40 
January 9 302 130 4 38 10 13 221 95 
February 9 75 27 - re - 9 15 27 
March 9 323 170 - - - 9 323 170 


4.15 days, n = 5) and 2000-2001 (11.40 + 3.63 days, Pine, and White Spruce, were used in proportion to 
n= 5). In 1999-2000, snow was generally packed or availability. Monocots, including Quackgrass (Elymus 
crusted with seeps and small streams remaining un- repens) and Timothy (Phleum pratense), were used less 
frozen. In 2000-2001, deep, powdery snow was com- _ than available, while dicots, including ash (Fraxinus 
mon, and most seeps were frozen by mid-winter. spp.), aster, Beaked Hazel, burdock, Choke Cherry 
(Prunus virginiana), Dogwood (Cornus stolonifera), 
Evening Primrose (Oenothera biennis), Fireweed (Epi- 
lobium angustifolium), goldenrod, Northern Wild Rai- 
sin (Viburnum cassinoides), raspberry (Rubus spp.), 
Soybean, Speckled Alder (Alnus incana), Swamp 
Thistle (Cirsium muticum), and willow (Salix spp.) 
were used in proportion to availability. 


Forage Selection 

Wild Turkeys foraged primarily on clovers (Trifo- 
lium spp.), asters (Aster spp.), goldenrods (Solidago 
spp.), and fertile fronds of the Sensitive Fern (Onoclea 
sensibilis) in the winter 1999-2000. Turkeys avoided 
species such as Cattail (Typha latifolia) and Meadow 
Sweet (Spiraea latifolia). Crops (n = 2) contained grass 
seeds and Sensitive Fern spore heads in late winter, Roost Site Characteristics 
confirming observations of turkeys feeding in the field. Trembling Aspen, American Basswood (Tilia ameri- 
Ostrich Fern (Matteuccia struthiopteris), Trembling cana), Jack Pine (Pinus banksiana), Eastern White 
Aspen (P. tremuloides) buds, Corn (Zea mays), Smooth Pine, White Spruce, and Eastern White Cedar (Thuja 
Wild Rose (Rosa blanda), and ragged moss (Brachy- occidentalis) were used as roost sites. Tree heights at 
thecium spp.) were other winter foods. Burdocks roosting sites averaged 13.9 + 0.8 m, and were higher 
(Arctium spp.), Sensitive Fern, and Soybeans (Glycine _ than the tree heights (8.8 + 0.7 m) measured at ran- 
max) comprised 21.4%, 23.0%, and 37.4% of the tur- dom sites (F, 5, = 20.98, P = 0.0001; Table 3). Mean 
keys’ diet, respectively, in 2000-2001. dbh of trees associated with roost sites, 37.7 + 3.0 cm, 

Forages used by turkeys differed from availability was significantly larger (F, 5, = 15.69, P = 0.0007) 
(c? = 40.38, df = 4, P < 0.0001; Table 2). Mosses _ than tree dbhs on random sites (24.1 + 1.7 cm). Roost 
were used in proportion to availability while ferns and __ site topographical elevations averaged 217.1 + 7.1 m 
allies, including Bracken (Pteridium aquilinum) and and were significantly higher (F, 5, = 4.49, P = 0.0455) 
Sensitive Ferns, were selected more than available. than the 188.8 + 11.3 m measured at random sites. Can- 
Conifer and allies, including Balsam Fir, Eastern White opy cover (73.3 + 8.8%), tree density (849.3 + 127.7 


TABLE 2. Chi-square analysis and Bonferroni Confidence Intervals for winter ground forage used by Eastern Wild Turkeys 
during winter in Noélville, near Sudbury, Ontario (v7 = 40.38, df = 4, P < 0.0001). 


Forage Forage Forage Proportion Proportion 90% Confidence 
Class Selected Available Selected Available Interval on Proportion 
Mosses 0.1 1.0 0.001 0.004 0.000-0.006° 
Ferns and allies 43.0 32.0 0.230 0.124 0.158-0.302° 
Conifers and allies 5.0 4.0 0.027 0.016 0.000-0.055° 
Monocots 0.1 13.0 0.001 0.050 0.000-0.006* 
Dicots 139.0 208.0 0.743 0.806 0.669-0.817° 
Total 187.2 258.0 


“Proportions greater than the upper confidence limit indicates use less than available. 
Proportions within the confidence limit indicates use equal to availability. 
‘Proportions less than the lower confidence limit indicates selection greater than available. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE 3. Comparison of physiographic characteristics of Eastern Wild Turkey roost (n = 12) and random (n = 12) sites in 


Noélville, near Sudbury, Ontario. 


Roost sites 


Habitat characteristic Mean SE 
Height (m) 13:9 0.8 
Tree dbh (cm) B07 3.0 
Canopy coverage (%) 133 8.8 
Density (trees/ha) 849.3 127 
Elevation (m) PAIN HA 
Distance to habitat edge (m) 97.5 Del 
Distance to open water (m) ABS 18.6 


trees/ha), distance to clearing (97.5 + 21.1 m), and dis- 
tance to open water (73.5 + 18.6 m) of roost trees did 
not differ (all P > 0.10) from those at random sites. 

Height of roost site was correlated with tree dbh 
(r= 0.78, n = 24, P = 0.003). There was no correlation 
between elevation and tree dbh (r = —0.36, n = 24, 
P = 0.245) nor elevation and tree height (7 = 0.04, n 
= 24, P = 0.900). Tree height was primarily responsi- 
ble for the differences between roosting and random 
sites, regardless of the covariate tree dbh (F, 5, = 3.45, 
P = 0.077). Tree dbh did not differ between roosting 
and random sites once the covariate height was statis- 
tically controlled (F, 5, = 15.87, P = 0.513). 


Discussion 

Winter mean home range sizes for turkeys released 
in northern Ontario (204 + 47 ha) were similar to 
those of turkeys introduced to Indiana (Miller et al. 
1985), and Iowa (Little and Varland 1981), but exceed- 
ed those reported in Ohio (Clark 1985) and in southern 
Ontario (Weaver 1989). Although previous studies 
reported a decline in monthly home range size be- 
tween November to March (Porter 1977; Miller et al. 
1985), this pattern was not observed in 1999-2000. 
This suggested that movement in late winter (1999- 
2000) was not limited by snow depth. On several occa- 
sions when snow covered local food resources, birds 
increased flight distance to otherwise unused parts of 
the home range. Snow depth was inversely related to 
winter and monthly home range size, confirming data 
from previous studies in Minnesota (Porter 1977), 
Indiana (Miller et al. 1985), Pennsylvania (Wunz and 
Hayden 1975), and southern Ontario (Weaver 1989). 

Glover and Bailey (1949) described the Wild Turkey 
as a nomadic feeder with a tendency to sample a wide 
variety of forages, primarily dictated by their avail- 
ability. The heavy use of ferns and allies was almost 
exclusively due to the selection of fertile fronds of 
the Sensitive Fern found in lowland hardwood sites 
and along seeps. Decker et al. (1991) reported that 
Sensitive Ferns were foraged in large quantities by 
Wild Turkeys because of their high nutritional content 
(crude protein 18.6%) and/or the high availability of 


Random sites 


Mean SE F P 
8.8 0.7 20.98 0.0001 
24.1 N57 15.69 0.0007 
70.4 9.7 0.05 0.8303 
764.3 110.9 0.25 0.6206 
188.8 ARS 4.49 0.0455 
64.8 7-7 1.42 0.2468 
122.1 34.7 152 0.2303 


this food in concentrated, relatively snow-free patches. 
Vander Haegen et al. (1989) reported that fields spread 
with manure were also important sources of winter 
food in Massachusetts, but the value of manure to win- 
tering turkeys in central Ontario was questionable, as 
emaciated birds were observed to ignore manure piles. 
Glover and Bailey (1949) reported wariness by turkeys 
of supplemental feeding sites with corn and oats in 
West Virginia when natural forage (e.g., wild grape, 
Vitis spp.) was absent. 

Standing crops, such as Soybean or clover, provid- 
ed nutritious food in moderate snow conditions during 
early winter. Soybeans contain protein, but also pro- 
duce trypsin inhibitors, which lower digestibility and 
fat absorption (McNaughton and Reece 1980, in Loesch 
and Kaminski 1989). Post-mortems in winter 2000- 
2001 (n = 5) verified that emaciated turkeys were 
under severe nutritional stress, and had catabolized 
significant amounts of muscle tissue. 

Forages that were present in the study area, but may 
have been underestimated in use, included conifer and 
hardwood trees. Although primarily ground feeders, 
Eastern Wild Turkeys are capable of flight, but forage 
obtained in the tree-canopy layer is difficult to sys- 
tematically record. Turkeys were observed feeding on 
Trembling Aspen buds on several occasions; however, 
the results indicate that prolonged periods of deep snow 
severely limited food availability for Wild Turkeys in 
northern Ontario. 

Roosting sites of turkeys were the tallest and largest 
trees, usually conifers, which were found at higher 
elevations. Conifers may reduce wind speed and heat 
loss (Kilpatrick et al. 1988). Tzilkowski (1971) found 
that turkey winter roosts were dominated by large decid- 
uous trees in Pennsylvania, suggesting that tree height 
and sturdiness were important factors. However, Cham- 
berlain et al. (2000) suggested that turkeys roost in the 
nearest suitable habitat at the end of the day. Although 
roost sites were closer to open water than random sites, 
this proximity appeared to be due to forage availability. 
Turkeys used snow to meet winter water requirements 
in this study. Exum et al. (1985) argued against Wild 
Turkey dependence on open water in southern Ala- 


1 
ii 


2004 


bama during spring and summer; however, Kilpatrick 
et al. (1988) suggested that winter foods in Rhode Is- 
land were too low in water content to meet the species’ 
needs. Roosting sites may have been closer to water 
because of the accelerated growth of trees associated 
with moist environments (Kilpatrick et al. 1988). 

The results of this study suggest that (1) winter food 
and roost site availability are the primary factors deter- 
mining successful turkey introductions in northern On- 
tario, (2) stands of Soybean or Corn are not sufficient 
to support Wild Turkey populations unless natural foods 
are available, and (3) snow depth is an important 
parameter influencing successful reintroduction of 
this species on the Precambrian Shield. 


Acknowledgments 

Financial and logistic support was provided by the 
West Arm/French River Wild Turkey Chapter, Ontario 
Federation of Anglers and Hunters, Cambrian College 
of Applied Arts and Technology, Laurentian University, 
Ontario Ministry of Natural Resources, Sudbury Dis- 
trict, and the Ontario Living Legacy Fund. We thank 
J. Craig and other landowners in Noélville for allow- 
ing fieldwork on their property, M. N. Hall for the 
Ontario Base Maps and Forest Resource Inventory 
data, and G. M. Courtin for meteorological data. 


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Received 24 March 2003 
Accepted 15 November 2004 


Fifteenth Census of Seabird Populations in the Sanctuaries of the 
North Shore of the Gulf of St. Lawrence, 1998-1999 


JEAN-FRANCOIS RAIL and GILLES CHAPDELAINE 
Canadian Wildlife Service, P. O. Box 10100, Ste-Foy, Québec G1V 4H5 Canada 


Rail, Jean-Francois, and Gilles Chapdelaine. 2004. Fifteenth census of seabird populations in the sanctuaries of the North 
Shore of the Gulf of St. Lawrence, 1998-1999. Canadian Field-Naturalist 118(2): 256-263. 


For the first time since the tradition began in 1925, the quinquennial census of seabirds in the Migratory Bird Sanctuaries of the 
North Shore of the Gulf of St. Lawrence was divided between two years in 1998-1999. Trends between 1993 and 1998-1999 
were variable across species and sanctuaries. In particular, following the large decreases in both species noted in 1993, the 
1998-1999 survey showed that Black-legged Kittiwakes had declined further, while the number of Herring Gulls had stabilized. 
Alcids were all doing well except for the Atlantic Puffin which showed severe yet unexplained drops at all major colonies. 
Law enforcement efforts appear reflected in seabird population trends, as well-patrolled sanctuaries such as St. Mary’s Islands 
seem to do well, whereas many species at the Baie des Loups and Ile a la Brume sanctuaries are far from their historical levels. 
In addition to a better law enforcement program, research is needed in order to identify other conservation problems that 
some species may be facing. 


Pour la premiere fois depuis 1925, l’inventaire quinquennal des oiseaux marins des refuges d’ oiseaux migrateurs de la Céte- 
Nord du Golfe Saint-Laurent fut mené sur deux ans, soit en 1998 et 1999. Les tendances des populations entre 1993 et 1998- 
1999 étaient variables selon |’espéce et le refuge. En particulier, suite a une décroissance notée chez les deux espéces en 1993, 
Vinventaire de 1998-1999 a démontré que la Mouette tridactyle avait encore diminué, tandis que le nombre de Goélands 
argentés s’ était stabilisé. Les populations d’ Alcidés ont toutes semblé en bonne santé sauf celle du Macareux moine dont les 
colonies importantes ont diminué de facon sévére et inexpliquée. Les efforts de protection par l’application de la loi se reflétent 
visiblement dans les tendances des populations d’ oiseaux de mer, puisque les refuges les mieux surveillés comme celui des 
Iles Sainte-Marie vont bien, alors que plusieurs espéces aux refuges de Baie des Loups et de l’ile 4 la Brume sont loin de leurs 
niveaux historiques. En plus d’un meilleur programme d’ application de la loi, la recherche est nécessaire pour identifier les 


autres problémes de conservation auxquels certaines espéces font face. 


Key Words: Seabirds, populations, sanctuaries, Gulf of St. Lawrence, Atlantic Puffin, larids, alcids. 


Apart from its major use in providing updated infor- 
mation on seabird numbers and distribution through- 
out the migratory bird sanctuaries on the North Shore of 
the Gulf of St. Lawrence (Figure 1), another major ap- 
plication of the results of this traditional survey is in 
identifying trends. Follow-up censuses have been car- 
ried out fairly regularly since the first census in 1925 
(Lewis 1925, 1931, 1937, 1942; Hewitt 1950; Tener 
1951; Lemieux 1956; Moisan 1962; Moisan and Fyfe 
1967; Nettleship and Lock 1973; Chapdelaine 1980, 
1995; Chapdelaine and Brousseau 1984, 1991), provid- 
ing the trends over a period of approximately 75 years, 
including the present survey. This survey also provides 
one of the ways we monitor the broader marine envi- 
ronment. In effect, these wide-ranging birds are acting 
as our sampling agents. As human impact on the seas 
increases, whether directly by exploitation of resources 
or marine pollution, or less directly by factors such as 
global warming, early indications of changes are vital. 
Clearly any changes in the size of seabird breeding col- 
onies are of major importance to conservationists to ori- 
ent any concrete action to be taken or to seabird biol- 
ogists to learn more about the regulation of seabird 
numbers and the wide range of factors known to influ- 
ence the birds. 


This article presents the current status of the 15 ma- 
rine bird species breeding in the sanctuaries in 1998- 
1999, and compares them with those of the 1993 census. 
Scientific names are given in Table 1. 


Methods 

The census of seabirds in the sanctuaries of the North 
Shore of the Gulf of St. Lawrence was conducted over 
two years. All of the technical details of this census 
(methods, estimates calculations, mapping of the col- 
onies, weather conditions) are discussed in Rail and 
Chapdelaine (2002) and summarized below for each 
family of birds. 

GaviiDs — We counted all Red-throated Loon nests by 
systematically walking around the ponds on the islands 
of each sanctuary. In a few exceptional cases we noted 
the presence of adults instead of landing on the island. 

HyDROBATIDS — We conducted a systematic count of 
active burrows. A burrow was considered active if we 
were able to reach an adult inside, or detect signs of 
recent occupation (soil freshly excavated or oily odour 
characteristic of petrels at the entrance of the burrow). 

ANATIDS — On Corossol Island, we used a system 
of quadrats from which we extrapolated and average 
density (nests/ha) for the entire area deemed suitable 


256 


RAIL and CHAPDELAINE: CENSUS OF SEABIRD POPULATIONS 


_/Me du Corossol —~Betchouane 


Gaspé Peninsula 


NEW BRUNSWICK 


ic Seaiiwiune Soran ogg “Gu, GF 
Watshishou 


lle a la Brume 


Gulf of St. Lawrence f~ = 4 


LABRADOR 57°00' W 


52°00' N 


QUEBEC 


yew Baie de Brador 
“ Saint-Augustin 
} ib Gros Mécatina 


\ 
t 


‘i Ce ony 2 oe lles Sainte-Mari 
) logging es Sainte-Marie, 


lles aux Perroquets 
Baie des Loups ‘ 


NEWFOUNDLAND 


ee 


FiGurE |. Location of the sanctuaries of the North Shore of the Gulf of St. Lawrence. 


for Common Eiders (Chapdelaine 1978*). In the Bet- 
chouane Bird Sanctuary, we conducted a systematic 
nest count on Calculot Island, and (as in preceding 
censuses) used a system of line-transects of varying 
length on Innu Island (see Caughley 1977). In sanc- 
tuaries featuring many islands (e.g., Watshishou, ile a 
la Brume, Baie des Loups), we counted all the nests 
on at least 27% of the land area and extrapolated an 
average density over the entire area of all the islands. 
On the Iles aux Perroquets and [les Sainte-Marie sanc- 
tuaries, extrapolation was done after systematic counts 
were made on most large islands, covering 57% and 
98% of total land area, respectively (but see Rail and 
Chapdelaine 2002 for details). 

PHALACROCORACIDS — Systematic counts of all Great 
and Double-crested Cormorant nests were carried out. 
At the Corossol Island colony, Double-crested Cormo- 
rant nests in treetops were counted from an elevated 
lookout point. 

Larips — In the large Herring Gull colony of Coros- 
sol Island, we sampled sub-colonies where the number 
of nests (Np) and the number of adults (Ni) were 
determined. Then, using the factor k (k=Np/Ni), we 
estimated the number of pairs in sectors where we 
counted only the number of adults present. In the 
Betchouane Sanctuary, all gull nests were counted sys- 
tematically, and were attributed to Herring and Great 
Black-backed gulls according to the observed propor- 
tions of adults of both species. In the other sanctu- 


aries all larids were censused with a combination of 

est counts and adult counts. Proportions of Arctic and 
Common terns in mixed tern colonies were not always 
determined, so the results for the two species are com- 
bined in Table 1. All Black-legged Kittiwake nests 
attended by adults were counted. 

ALCIDS — We carried out systematic counts of eggs 
and active burrows in colonies of Atlantic Puffins, 
Razorbills and Common Murres that could be accessed 
with minimal disturbance. Elsewhere we counted indi- 
viduals at the colony and on adjacent waters from a 
distance. All colonies with large numbers of Common 
Murres were carefully avoided because of their sensi- 
tivity to disturbance. In particular, at the Iles Sainte- 
Marie Sanctuary, where 87% of breeding Common 
Murres on the North Shore are found, 98% of the 
population estimate for the Common Murre results 
from adult counts. For Atlantic Puffins nesting on the 
Blacklands Island in the Baie des Loups Sanctuary, we 
used the factor k method described above for larids. 
On Ile aux Perroquets in the Baie de Brador Sanctuary, 
we used evenly spaced grids and line transects to de- 
limit the area of both the puffin and Razorbill colonies, 
evaluate mean nest densities, and then extrapolated 
the populations (see Nettleship 1973* ; Chapdelaine 
1978*). The Black Guillemot population was estimated 
from adult bird counts around the islands, although 
exceptionally we recorded a pair when a nest was 
found with no adult in sight. 


Vol. 118 


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2004 RaiL and CHAPDELAINE: CENSUS OF SEABIRD POPULATIONS 259 
TABLE 2. Changes in the numbers of seabirds in sanctuaries on the North Shore of the Gulf of St Lawrence, Québec, 1988 to 
1998-1999. 

Years of survey Compound annual growth* 

rate by period 

Species 1988 1993 1998-1999 1988-1993 1993-1998** 
Red-throated Loon 76 66 80 -2.82% 3.21% 
Leach’s Storm-Petrel 1744 1840 718 1.07% -18.78% 
Great Cormorant 86 78 340 -1.95% 24.54% 
Double-crested Cormorant 4558 3472 2830 -5.44% -3.49% 
Common Eider 8536 14548 13072 10.66% -2.10% 
Ring-billed Gull 288 104 484 -20.37% 31.04% 
Herring Gull 16195 4879 4929 -24.00% 0.49% 
Great Black-backed Gull 1883 2284 2381 3.86% 1.15% 
Black-legged Kittiwake 8536 6294 3816 -6.09% -10.02% 
Caspian Tern 15 0 0 -100.00% - 
Common and Arctic Terns 1350 545 294 -18.14% -12.03% 
Common Murre 26049 30829 30111 S370 -0.39% 
Razorbill 7036 8389 13953 35200 8.58% 
Black Guillemot 521 411 651 -4.74% 8.70 % 
Atlantic Puffin 35142 46684 29030 5.68% -7.92% 


log. N (t) - log. N (0) 
so — 100% 
t 


** Populations in sanctuaries censused in 1999 were back-calculated to 1998 to get the compound annual growth rate for 


all North Shore sanctuaries after 5 years 


Results 

In the eight sanctuaries censused in 1993 and 1998- 
1999 (Gros Mecatina excluded), the total number of 
birds had decreased by 15% or 17 734 individuals (see 
Table 1 for details). This is mostly due to the dra- 
matic 38% decline of the Atlantic Puffin (-17 654 indi- 
viduals), which was by far the most abundant seabird 
in the North Shore sanctuaries in 1993 (but was out- 
numbered by the Common Murre in 1999). Leach’s 
Storm-Petrels, Black-legged Kittiwakes and Common 
and Arctic Terns also declined at a very rapid rate 
between 1993 and 1998-1999 (Table 2). Double-crested 
Cormorants numbers also diminished. The Razorbill is 
the only abundant species showing a large increase, 
with the 1998-1999 population estimate exceeding by 
5564 individuals the result of 1993 (+66%). This species 
has made a spectacular comeback since 1972, when less 
than 3000 individuals were recorded in the sanctuaries 
(compared to 14 341 in 1998-1999). The small popula- 
tions of Great Cormorants, Ring-billed Gulls and Black 
Guillemots also grew rapidly between 1993 and 1998- 
1999, as did Red-throated Loons but more slowly. In 
four species, numbers appeared rather stabilized, the 
Common Eider and Herring Gull after having showed 
very sharp trends between 1988 and 1993 (annual 
growth rates of +11% and -24%, respectively), the 
Great Black-backed Gull and Common Murre follow- 
ing less striking but significant rises (annual growth 


rate of +3.9 and +3.4%, respectively). Finally, no Cas- 
pian Terns were found for the second quinquennial 
census in a row. 

We visited the Corossol Island Sanctuary on 30 and 
31 May 1998. Major increases in large gulls (Herring 
and Great Black-backed gulls) were noted. Populations 
of alcids also expanded, and in particular the number 
of Black Guillemots more than doubled compared to 
1993. Puffins were recorded for the first time during 
quinquennial censuses, bringing up to 10 the number 
of seabird species breeding there. The Common Eider 
was also more abundant. On the other hand, the colony 
of Leach’s Storm-Petrels was reduced to less than half 
between 1993 and 1998, and Black-legged Kittiwakes 
also declined quite rapidly. Double-crested Cormorants 
were down 25%, and colony localization had seemed 
to change a bit, possibly prompted by forest decay 
caused by their long-term occupation (nests are in tree- 
tops of conifers) in some areas. 

On 2 and 3 June 1998, we visited the Betchouane 
Bird Sanctuary. The most important change in the sea- 
bird community was the 36% decrease in Common 
Eider, by far the most abundant bird there. The Razor- 
bill apparently continued its spectacular comeback, 
with a 72% population increase between 1993 and 
1998. Herring Gulls, Great Black-backed Gulls, Black- 
legged Kittiwakes and Atlantic Puffins were almost 
stable or slightly increasing. 


260 


After a tremendous population explosion following 
1982, the population growth of the Common Eider in 
Watshishou appeared to have slowed down consid- 
erably between 1993 and 1998. The Black Guillemot 
population increased threefold during the same period. 
Double-crested Cormorant and Great Black-backed 
Gull numbers showed opposite trends; the former spe- 
cies underwent a 50% increase whereas the latter was 
reduced by half. Herring Gull population size appeared 
slightly reduced compared to 1993, but the 1998 result 
was by far the highest for the species in Watshishou, 
except for 1993. The 1998 survey confirmed a cata- 
strophic (-96%) and inexplicable decline of terns in 
Watshishou since 1988. In the past, this sanctuary, con- 
sisting of more than 200 islands, always had a fair 
number of terns (range 128 — 1490 individuals), and 
was the most important sanctuary on the North Shore 
in that regard. In 1998, however, despite better cover- 
age (more islands were visited than in previous cen- 
suses), we found only 19 breeding pairs distributed 
in 5 very small colonies. The Watshishou Bird Sanc- 
tuary was visited on 6, 7 and 8 June 1998. 

The following year (in 1999), seabirds were cen- 
sused on 14 June at the Ile 4 la Brume Sanctuary. The 
Common Eider had decreased slightly since 1993, 
but was still relatively abundant. Population trends 
were up for Herring and Ring-billed gulls; however 
numbers of these two species are rather small and 
tended to vary a lot in the past. The number of Great 
Black-backed Gulls dropped further between 1993 
and 1999, reaching its lowest level ever in 1999. Terns 
declined sharply during the same period, and in fact 
since 1988 their population trends at the Ile a la Brume 
(-84%) and Watshishou sanctuaries are similar. Status 
of the very small populations of Red-throated Loon 
and Razorbill are still precarious, as their numbers 
remained stable between 1993 and 1999 at two and 
eight individuals, respectively. Finally, no Caspian 
Terns were found for the second quinquennial census 
in a row at Ile a la Brume, the only traditional breed- 
ing site for this species in the Province of Québec. 

We visited the Baie des Loups Bird Sanctuary on 
18 June. The most abundant bird there, the Atlantic 
Puffin, had declined by 40% since 1993. The Great 
Black-backed Gull population also decreased notice- 
ably (36%). Breeding Leach’s Storm-Petrels apparent- 
ly deserted the sanctuary, which leaves the Corossol 
Island Sanctuary as the only active breeding site of that 
species on the North Shore, and maybe in the whole 
province. On the brighter side, the Common Eider pop- 
ulation increased by 50%. Also, the Common Murre 
and the Razorbill, once abundant in the sanctuary but 
which had declined steadily up to the very low and 
worrying levels observed in 1993, bounced back and 
increased threefold between 1993 and 1999. Red- 
throated Loon, Herring Gull, terns, and Black Guille- 
mot numbers remained fairly stable. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


The Iles aux Perroquets Bird Sanctuary was visited 
on 11 and 21 June 1999. The population of Atlantic 
Puffin showed a similar drop (-53%) as in the other 
large colonies on the North Shore. In contrast to the 
general upward trend observed in these species at the 
other sanctuaries, a decrease in Common Murres and 
Razorbills was noted at the Iles aux Perroquets Bird 
Sanctuary in 1999. The number of Common Eiders 
declined only slightly. Except for the Black-legged 
Kittiwakes whose population collapsed (-82%) between 
1993 and 1999, numbers of larids were quite stable. 
With only 11 nests found in 1999, the kittiwake may 
well disappear as a breeding species at the Iles aux 
Perroquets in the future. On the positive side, the two 
smallest seabird populations breeding in the sanctuary 
in 1993, those of the Red-throated Loon and Black 
Guillemot, both showed large increases (140% and 
164%, respectively). Also worthy of notice is the addi- 
tion of the Great Cormorant as a breeding species in 
the sanctuary. 

The Iles Sainte-Marie Bird Sanctuary (visited on 
10, 11, 13, 17 and 19 June) was again found to have 
the highest seabird diversity (12 breeding species) 
and abundance (total of 39 316 individuals). Common 
Murres represented two-thirds of the seabird popula- 
tion with an estimated 26 156 breeding birds. The 
number of murres appeared quite stable between 1993 
and 1999, after the spectacular and steady recovery 
of the species following 1972 (when only 4120 birds 
were counted). Razorbills show the same long-term 
trend as murres, but were still going up in 1999 (116% 
rise compared to 1993). The Great Cormorant colony 
expanded unexpectedly (+318%) to become the largest 
of this species on the North Shore. Increases were noted 
in Black-legged Kittiwakes, Black Guillemots and terns 
as well. Numbers of Double-crested Cormorants, Com- 
mon Eiders, and Atlantic Puffins were reduced rough- 
ly by half. Herring Gulls were again found to be declin- 
ing very sharply, whereas Great Black-backed Gulls 
showed a moderate decrease. Finally, the number of 
Red-throated Loons in 1999 was only one pair short of 
the highest counts which were recorded on the two pre- 
ceding censuses. 

A little farther to the east, a few small offshore 
islands constitute the Gros Mécatina Bird Sanctuary, 
which was created in 1996. On 23 June 1999, the sanc- 
tuary was included for the first time in a quinquennial 
census. Species diversity was impressive for such a 
small sanctuary, as nine seabird species were found 
breeding. Numbers of each species were low though, 
and the most abundant species was the Razorbill with 
388 birds observed. According to naturalists such as 
Coues (1862), these islands were once inhabited by tens 
of thousands of Common Murres, so the sanctuary has 
definite potential for seabird conservation if it is well 
protected. 


2004 


The Baie de Brador Bird Sanctuary is well-known 
for its colonies of Atlantic Puffin and Razorbill. On 28 
and 29 June 1999 however, we were quite surprised 
to discover the addition of Black-legged Kittiwakes 
and terns breeding there for the first time, as well as 
Common Murres which had not bred there since 
1965. Herring Gulls and Great Black-backed Gulls 
seemed well established as breeders for the second 
census in a row (they were present in very small num- 
bers or absent prior to 1993). Also, as in most other 
sanctuaries, Razorbill numbers grew very fast between 
1993 and 1999 (+175%). Nevertheless, all these results 
were darkened by the finding that Atlantic Puffins had 
declined by a third since 1993, when the population 
seemed on its way to get back to historic levels. 
Because it has always been the stronghold of puffins 
in our province, the Baie de Brador Bird Sanctuary 
plays a leading role in the conservation of the species 
in Québec. 


Discussion 

Total number of seabirds recorded in 1998-1999 in 
the North Shore Bird Sanctuaries dropped 15% (or 
17 734 birds, Gros Mécatina excluded). But most of 
this decrease could be attributed to the Atlantic Puffin 
whose 38% decline represent a loss of 17551 birds. 
Population trends in other species were variable, as 
four species increased rapidly, four species decreased 
at a fast rate, and the others showed more stable or 
moderate trends. Numbers of Razorbills and Black 
Guillemots increased at a very rapid rate in almost 
every sanctuary (where they are present). The only Great 
Cormorant colony recorded during the 1993 census was 
found at the [les Sainte-Marie Bird Sanctuary; by 
1999, that colony had grown so much that immigration 
must be partly involved. Also, three new colonies of 
this species were found elsewhere in the sanctuaries 
in 1999. The Ring-billed Gull is the last seabird species 
which increased at a very fast rate since 1993. How- 
ever, this species is poorly represented in the North 
Shore Bird Sanctuaries, where their numbers always 
fluctuated widely in the past. Instead of suggesting a 
real population trend, our results may rather reflect 
colony displacements as the species exhibits poor site 
fidelity on the North Shore. 

The Atlantic Puffin was not the only species show- 
ing a sharp decline between the last two censuses of 
the North Shore sanctuaries. The case of the Leach’s 
Storm-Petrel is equally worrying; the species was found 
breeding in four sanctuaries in 1988, but in 1998- 
1999 only the Corossol Island colony was still active, 
and its number of nests was reduced to less than half 
compared to 1993. Moreover, this may be the only 
Leach’s Storm-Petrel colony left in the whole province. 
Thus the status of the species appears precarious in 
Québec. The Black-legged Kittiwake was found breed- 
ing in six sanctuaries in 1998-1999, whereas in the past 
it had been breeding in no more than four sanctuaries. 


RAIL and CHAPDELAINE: CENSUS OF SEABIRD POPULATIONS 


261 


However, over 90% of the kittiwakes in the sanctu- 
aries are concentrated at the Corossol Island colony, 
where a 39% drop was observed between 1993 and 
1998. In fact the colony decreased by 55% since 1988 
and the trend was even faster in the last five years. 
Observations made on diet and breeding success of 
larids at Corossol Island in 1996-1998 showed that 
kittiwake breeding success was severely affected by 
Herring Gull predation on large chicks, especially when 
capelin appeared less available. The poor kittiwake 
productivity measured in 1997-1998 (0.60 and 0.27 
young fledged per pair, respectively) means that re- 
cruitment is likely not enough to compensate for nat- 
ural mortality, so that further decline of the colony is 
expected. Terns, after the fashion of Ring-billed Gulls, 
are poorly represented in the sanctuaries and their 
population trends partly reflect the unpredictability 
of their breeding activities. However, their numbers 
dropped severely for a second quinquennial census in 
a row, resulting in a 80% decline since 1988. The case 
of the Watshishou Bird Sanctuary, which has always 
been the most important for terns, is particularly intrigu- 
ing as that tern population reached an unprecedented 
low level in 1998. 

Among species with near-stable population trends 
between 1993 and 1998-1999 are the Common Eider 
and the Common Murre, two species whose numbers 
finally appeared to level off after spectacular increases 
were noted in each of the four preceding censuses. 
Herring Gull population trends on the North Shore 
(and especially at Corossol Island) have been corre- 
lated with cod fishery, an industry that provided signif- 
icant amounts of fish offal available to gulls (Chap- 
delaine and Rail 1997). After a boom in the early 1980s, 
cod fishery and Herring Gull populations collapsed 
concurrently in the late 1980s-early 1990s. But in 1998- 
1999 it seemed that after a drastic (-78%) decline be- 
tween 1988 and 1993, the Herring Gull population was 
stabilized, perhaps to a more “natural” level. The num- 
ber of Red-throated Loons increased only slightly be- 
tween the last two surveys, yet the 1998-1999 total 
(82 individuals) is the highest ever recorded in the 
sanctuaries. The Double-crested Cormorant population 
declined but only at a slow rate after 1993, and in 1998- 
1999 the total population estimate for the sanctuaries 
was still the third highest since 1925. The steady rise 
of Great Black-backed Gull numbers continued to slow 
down to be nearly stable between 1993 and 1998. Final- 
ly, the Caspian Tern did not breed in 1993 and 1999 at 
the fle 4 la Brume Sanctuary, where it consistently nest- 
ed between 1925 and 1988. The more time passes by, 
the more unlikely this species will be back, unless 
appropriate measures are taken to prevent disturbance 
of the only breeding site of Caspian Terns in Québec. 

While some population trends varied regionally, 
others suggest large-scale factors are involved. In 
particular, Black-legged Kittiwake and Herring Gull 
declines since the end of the 1980s are reported not only 


262 


in North Shore sanctuaries, but also at all large colo- 
nies around the Gaspé Peninsula and at the Magdalen 
Islands (Chapdelaine et al. 2003*). Atlantic Puffin num- 
bers dropped at all major North Shore concentrations 
between 1993 and 1999. On the other hand, Razorbill 
colonies continued to expand everywhere in the Gulf 
and in the Estuary (Chapdelaine et al. 2001), and the 
small populations of Great Cormorant on the North 
Shore and around the Gaspé Peninsula grew signifi- 
cantly since 1988-1990. So far we have presented some 
evidence that the Herring Gull population had been 
influenced by cod fishery activities (Chapdelaine and 
Rail 1997), and that an abundance of small fish such 
as sandlance (Ammodytes sp.) and Capelin (Mallotus 
villosus) had a positive effect on alcid breeding per- 
formance and populations (Chapdelaine and Brousseau 
1991, 1996; Rail et al. 1996). But most population 
trends are left unexplained. 

The fact that law enforcement efforts have not been 
distributed uniformly across sanctuaries probably ex- 
plains why, in sanctuaries such as Ile a la Brume and 
Baie des Loups, many seabird populations are at low 
levels and signs of disturbance are omnipresent. Con- 
versely, the well-patrolled [es Sainte-Marie Bird 
Sanctuary appears in good shape. Seabird conservation 
in North Shore bird sanctuaries face many challenges: 
ecotourism, disturbance, poaching, episodic fox inva- 
sion in the sanctuaries, seabird bycatch in nearby fish- 
ing nets, along with ecological factors at all scales. This 
stresses the need to maintain and improve law enforce- 
ment and public educational programs, and highlights 
the importance of research to tackle the factors respon- 
sible for the observed seabird population trends. 


Acknowledgments 

Many thanks to all those who contributed to this. 
Fieldwork was carried out with the help of our col- 
leagues Pierre Brousseau and Jocelyn Thibault, Gilles 
Falardeau with his assistants Linda Burr and Daniel 
Daigneault, and Canadian Wildlife Service wardens 
Freddy Strickland and Wilson Evans. Members of Parks 
Canada in Mingan and assisting volunteers also gave 
us a big hand, as well as Andrew Rowsell from Fish- 
eries and Oceans Canada in Blanc-Sablon. Finally, 
our special thaughts go to M. Gallienne and his family 
for their hospitality on Corossol Island. 


Documents Cited (marked * in text) 

Chapdelaine, G. 1978. Onziéme inventaire des oiseaux colo- 
niaux des refuges de la c6te nord du Golfe Saint-Laurent 
et révision globale de la fluctuation des populations depuis 
1925 jusqu’a 1977. Service canadien de la faune, région 
du Québec, Environnement Canada, Sainte-Foy, 166 pages. 

Chapdelaine, G., P. Brousseau, and J.-F. Rail. 2003. 
Banque informatisée des oiseaux marins du Québec 
(BIOMQ). Database. Last update 12 March 2003. Environ- 
ment Canada, Canadian Wildlife Service, Québec Region. 

Nettleship, D. N. 1973. Census of seabirds in the sanctuaries 
of the North Shore of the Gulf of St. Lawrence, summer 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


1972. Studies on northern seabirds No 20. Canadian Wild- 
life Service Report, Ottawa, 160 pages. 


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Chapdelaine, G. 1980. Onziéme inventaire et analyse des 
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Chapdelaine, G. 1995. Fourteenth census of seabird popu- 
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Chapdelaine, G., and P. Brousseau. 1984. Douzieme inven- 
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Chapdelaine, G., and P. Brousseau. 1991. Thirteenth cen- 
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Chapdelaine, G., and P. Brousseau. 1996. Diet of Razor- 
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Chapdelaine, G., A. W. Diamond, R. Elliot, and G. J. 
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Received 16 June 2003 
Accepted 5 July 2004 


Notes 


Observations of Above-Surface Littoral Foraging in Two Sea Ducks, 
Barrow’s Goldeneye, Bucephala islandica, and Surf Scoter, Melanitta 
perspicillata, in Coastal Southwestern British Columbia 


DEBORAH L. LACROIX!, KENNETH G. WRIGHT?, and DANIEL KENT? 


'Centre for Wildlife Ecology, Simon Fraser University, Burnaby, British Columbia VSA 1S6 Canada 
26090 Blink Bonnie Road, West Vancouver, British Columbia V7W 1V8 Canada 
>Vancouver Aquarium Marine Science Centre, Vancouver, British Columbia V6B 3X8 Canada 


Lacroix, Deborah L., Kenneth G. Wright, and Daniel Kent. 2004. Observation of above-surface littoral foraging in two sea 
ducks, Barrow’s Goldeneye, Bucephala islandica, and Surf Scoter, Melanitta perspicillata, in coastal southwestern 
British Columbia. Canadian Field-Naturalist 118(2): 264-265. 


Barrow’s Goldeneyes (Bucephala islandica) and Surf Scoters (Melanitta perspicillata) were observed on four separate occa- 
sions, by three different observers, foraging on Bay Mussels (Mytilus trossulus) above the water surface. This unique foraging 
behaviour could be attributed to diurnal spring tides and reduced lower intertidal mussel abundance. 


Key Words: Barrow’s Goldeneye, Bucephala islandica, Surf Scoter, Melanitta perspicillata, foraging behaviour, sea ducks, 


Bay Mussels, Mytilus trossulus, British Columbia. 


The sea ducks, Barrow’s Goldeneye (Bucephala 
islandica) and Surf Scoter (Melanitta perspicillata), 
winter in abundance along the coast of British Colum- 
bia where they forage diurnally on marine inverte- 
brates in bays, harbours, beaches, and inlets (Bellrose 
1980; Vermeer 1981, 1982; Vermeer and Bourne 1984; 
Campbell et al. 1990; Savard et al. 1998; Eadie et al. 
2000). In rocky habitats, both sea ducks forage pre- 
dominantly on Bay Mussels (Mytilus trossulus) (for- 
merly classified as M. edulis; McDonald and Koehn 
1988, McDonald et al. 1991) (Vermeer and Levings 
1977; Hirsch 1980; Vermeer 1981, 1982; Vermeer and 
Bourne 1984; Lacroix 2001). Both sea ducks dive to 
locate and retrieve mussels. Mussels are pried from the 
substrate and swallowed whole (Savard et al. 1998; 
Eadie et al. 2000). 

Three observers, on four separate occasions, ob- 
served Surf Scoters and/or Barrow’s Goldeneyes for- 
aging on Bay Mussels above the water surface in coastal 
southwestern British Columbia. The first observation, 
on 22 February 1999, involved juvenile male and fe- 
male Surf Scoters and Barrow’s Goldeneyes eating 
exposed mussels at a breakwater on Popham Island. 
The event was photo-documented by D. Kent. On the 
second occasion, on 7 March 1999, 1 juvenile male and 
7 females, mostly juveniles, Barrow’s Goldeneyes and 
3 Surf Scoters, (1 female and 2 immature males), re- 
moved mussels from a rocky point at Cape Roger 
Curtis, while they sat on a reef, 1.5 m above the water 
line. Later that day a flock of approximately 12 Bar- 
row’s Goldeneyes, mostly juveniles, was seen feeding on 
mussels while hauled out on a reef on Hermit Island. 
This above-surface foraging tactic was also noticed 


later in March when three juvenile Surf Scoters, and 
six Barrow’s Goldeneyes, mostly juveniles, fed on ex- 
posed mussels on large boulders. All the observations 
coincided with low diurnal tides. The tide height ranged 
between 1.74 to 2.16 m, above Chart Datum (a.c.d.). 
We are unaware of any previously published or unpub- 
lished accounts of this foraging behaviour. Our multiple 
observations suggest that the behaviour occurs frequent- 
ly but has gone unreported. 

All of these observations were made during late 
February and March in Howe Sound and Burrard 
Inlet, British Columbia, located in the same geograph- 
ic vicinity (49°19.32’N, 123°09.92’W — 49°21.60’N, 
123°29.15’W). These observations share several sim- 
ilar characteristics including: (1) all ducks were feed- 
ing on Bay Mussels; (2) the observations were made 
during low diurnal tides; (3) only small groups, often 
consisting of mixed-species flocks, were exhibiting this 
foraging behaviour, and (4) these foraging groups 
consisted mostly of juveniles. 


Discussion 

Bay Mussels are a dominant species in protected 
coastal rocky intertidal areas (Seed and Suchanek 
1992: Ricketts et al. 1995). In our observation area, 
the vertical distribution of the Bay Mussel ranges from 
1.5 to 3.7 m, a.c.d. (Quayle 1978). Although Bay Mus- 
sels are intertidal, mussel beds are rarely completely 
exposed during daylight hours in winter as the lowest 
low tides of the semidiurnal tidal regime occur at night 
(Thomson 1981). As winter advances into spring, the 
lowest low tides are diurnal; therefore expose mussel 
beds during daylight hours (Figure 1). Exposed mussels 


264 


2004 


FIGURE | 


Percent of daylight hours 
fully submerged 


Oct Nov Dec Jan Feb Mar Apr May 


Time 


Ficure |. The percent of daylight hours that Bay Mussel beds 
are completely submerged in coastal British Columbia. 
Mussel submergence time was estimated by calculat- 
ing the number of hours the tide was above 3.7 m, 
a.c.d, using Canadian tide charts (1999), during day- 
light hours, between sunrise and sunset. 


may be more vulnerable to diving ducks such as Surf 
Scoter and Barrow’s Goldeneye as the ducks do not 
need to expend energy diving to reach the mussels. 
However, Smeathers and Vincent (1979) found that 
mussels exposed to air have twice the tensile strength 
of those submerged. If the exposed mussels have un- 
dergone some degree of desiccation, they may require 
more energy to remove than submerged ones, and 
may therefore, not be as profitable as first postulated. 
Alternatively, juvenile and sub-adult Surf Scoters and 
Barrow’s Goldeneyes may be forced to feed on exposed 
mussels owing to the reduced mussel abundance and 
distribution from over-winter predation. The lower por- 
tion of the distribution of Bay Mussels is determined 
by biological factors, primarily predation from the 
Ochre Sea Star (Pisaster ochraceus) (Seed and Sucha- 
nek 1992; Quayle 1978) and sea ducks (Lacroix 2001). 
Through the winter, the combination of sea star and sea 
duck predation may eliminate the lower distribution 
of mussels, hence reducing their overall abundance. 
It is therefore plausible that the ducks observed were 
forced to forage on the less profitable prey (1.e., the ex- 
posed mussels) because there are few or no submerged 
mussels in the lower portion of their distribution. 


Acknowledgments 

We thank R. W. Butler and A. J. Erskine for making 
valuable suggestions to the manuscript. D. L. Lacroix 
received funding from a postgraduate NSERC schol- 
arship and from the Canadian Wildlife Service Geor- 
gia Basin Initiative. 


Literature Cited 

Bellrose, F. C. 1980. Ducks, geese and swans of North Amer- 
ica. Stackpole Books, Harrisburg, Pennsylvania. 540 pages. 

Campbell, R. W., N. K. Dawe, I. McTaggart-Cowan, J. 
M. Cooper, and G. W. Kaiser. 1990. The birds of British 


NOTES 


265 


Columbia. Volume |. Nonpasserines, introduction and 
loons through waterfowl. Royal British Columbia Museum, 
Victoria, British Columbia. 513 pages. 

Canadian Hydrographic Service, Fisheries and Oceans 
Canada. 1999. Canadian tide and current tables. Volume 
5. Minister of Fisheries and Oceans Canada. 120 pages. 

Eadie, J. M., J.-P. Savard, and M. L. Mallory. 2000. 
Barrow’s Goldeneye (Bucephala islandica). Pages 1-32 
in The Birds of North America. (548). Edited by A. Poole 
and F. Gill. The Academy of Natural Sciences, Philadelphia, 
Pennsylvania and the American Ornithologists’ Union, 
Washington, D.C. 

Hirsch, K. V. 1980. Winter ecology of sea ducks in the inland 
marine waters of Washington. M.Sc. thesis, University of 
Washington, Seattle, Washington. 92 pages 

Lacroix, D. L. 2001. Foraging impacts and patterns of win- 
tering surf scoters feeding on bay mussels in coastal Strait 
of Georgia, British Columbia. M.Sc. thesis, Simon Fraser 
University, Burnaby, British Columbia. 126 pages. 

McDonald, J. H., and R. K. Koehn. 1988. The mussels 
Mytilus galloprovincialis and M. trossulus on the Pacific 
coast of North America. Marine Biology 99: 111-118. 

McDonald, J. H., R. Seed, and R. K. Koehn. 1991. Allo- 
zymes and morphometric characters of three species of 
Mytilus in the Northern and Southern Hemispheres. Marine 
Biology 111: 323-333. 

Quayle, D. B. 1978. A preliminary report on the possibili- 
ties of mussel culture in British Columbia. Fisheries and 
Marine Service Technical Report 815. 37 pages. 

Ricketts, E. F., J. Calvin, J. W. Hedgpeth, and D. W. Philips. 
1995. Between pacific tides. Fifth Edition. Stanford Uni- 
versity Press, Stanford, California. 

Savard, J-P. L., D. Bordage, and A. Reed. 1998. Surf Scoter 
(Melanitta perspicillata). Pages 1-28 in The Birds of North 
America. (363) Edited by A. Poole and F. Gill. The Aca- 
demy of Natural Sciences, Philadelphia, Pennsylvania and 
the American Ornithologists’ Union, Washington, D.C. 

Seed, R., and T. H. Suchanek. 1992. Population and com- 
munity ecology of Mytilus. Pages 87-157 in The mussel 
Mytilus: ecology, physiology, genetics and culture. Edited 
by E. Gosling. Developments in Aquaculture and Fisher- 
ies Science 25, Elsevier, Amsterdam. 589 pages. 

Smeathers, J. B., and J. F. Vincent. 1979. Mechanical prop- 
erties of mussel byssus threads. Journal of Molluscan 
Studies 45: 219-230. 

Thomson, R. E. 1981. Oceanography of the British Columbia 
coast. Canadian Special Publication Fisheries and Aquatic 
Science 56. 291 pages. 

Vermeer, K. 1981. Food and populations of Surf Scoters in 
British Columbia. Wildfowl] 32: 107-116. 

Vermeer, K. 1982. Food and distribution of three Bucephala 
species in British Columbia. Wildfowl 33: 22-30. 

Vermeer, K., and N. Bourne. 1984. The White-winged Scoter 
diet in British Columbia waters: resource partitioning with 
other scoters. Pages 62-73 in Marine birds: their feeding 
ecology and commercial fisheries relationships. Edited by 
D. N. Nettleship, G. A. Sanger, and P. F. Springer. Pro- 
ceedings of the Pacific Seabird Group Symposium, Seattle, 
Washington, 1982. 

Vermeer, K., and C. D. Levings. 1977. Populations, biomass 
and food habits of ducks on the Fraser Delta intertidal 
area, British Columbia. Wildfowl 28: 49-60. 


Received 23 July 2001 
Accepted 6 August 2004 


266 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


New Records of Cyperaceae and Juncaceae from the Yukon Territory 


Stuart A. HARRIS! AND PETER W. BALL? 


'Department of Geography, University of Calgary, Calgary, Alberta T3A 1E4 Canada 
*Department of Botany, Erindale College, University of Toronto in Mississauga, Mississauga, Ontario LSL 1C6 Canada 


Harris, Stuart A., and Peter W. Ball. 2004. New records of Cyperaceae and Juncaceae from the Yukon Territory. Canadian 


Field-Naturalist 118(2): 269-270. 


Two new species of Cyperaceae are reported, viz., Carex hoodii, and Eleocharis elliptica. Also, range extensions for eight 


species of Carex, Eriophorum, and Juncus are listed. 


Key Words: Vascular plants, Yukon Territory, new records, range extensions. 


In the last ten years, considerable advances have 
taken place in the knowledge of the vascular flora of 
the Yukon Territory (Cody, 1996, 2000; Cody et al., 
1998, 2000, 2001, 2002, 2003). In spite of this, the 
flora is still incompletely known, as indicated by the 
frequent and substantial additions. 

In 1980, the first author commenced detailed field 
studies of the distribution of permafrost in the south 
and central Yukon Territory, together with the nature 
and dynamics of the associated landforms (Harris 
1998, 2004; see also Wall et al. 1987). As part of the 
work, plants were collected and have been system- 
atically identified. For the more difficult groups, the 
material was sent to specialists who have helped in 
the identification. P.W.B. carried out the task of iden- 
tifying the Cyperaceae and some of the Juncaceae. 
The result is a substantial collection of sheets that are 
in the collection of the first author (UAC #60 000 — 
74 000). Comparison of these with the published lists 
of species and their distributions indicates that there 
are at least two species not previously described for 
the Yukon Territory, as well as some range extensions 
and name changes of varying importance. This paper 
will describe these changes. The nomenclature used 
follows that in the recent Flora of North America, 
volume 23 (Flora of North America Editorial Com- 
mittee 2002). 


Species New to the Yukon 

CYPERACEAE 

Carex Hoodii Boott — YUKON. Hart River road at 
1180 m elevation, east of the Dempster Highway at 
km 78, about 138°14'W., 64°30'N. (UAC 66735). 

It occurs on the wet shrub tundra between the clumps of 
Betula glandulosa and Salix spp., and is unlike any of the 
other Carex species. This species should be added to the list 
of rare plants in the Yukon (Douglas et al. 1981). 


Eleocharis elliptica Kunth. — YUKON. Klondike River 
bridge, Dempster Highway at 138°44'W., 63°59'N. 
(UAC 70230). 

This North American species is widespread in Alberta 
(Moss 1983, page 158), and should be looked for elsewhere 
in the southern Yukon Territory. It is characterised by yellow, 
orange or brown achenes with 12-20 vertical ridges. It should 
be added to the list of rare plants of the Yukon Territory 
(Douglas et al. 1981). 


Range Extensions in the Yukon 

CYPERACEAE 

Carex foenea Willdenow — YUKON. Tagish Camp- 
ground, 134°15'W., 60°19'N. (UAC 70310), and Lucky 
Lake, east of Watson Lake, 134°30'W., 60°O1'N. (UAC 
61656). 

It is probably present at scattered localities across the south- 
ern Yukon Territory in wet places in Lodgepole Pine forests. 
It has previously been found in three localities (Cody, 1994, 
page 149: Cody et al. 1998, page 301) including Watson Lake. 


Carex glareosa Wahlenburg — YUKON. Minto Landing 
in grassy glades at 136°53'W., 62°03'N. (UAC 70268). 

Douglas et al. (1981), and Cody (1994, page 440; 1996; 
Cody et al. 1998, page 301) have previously reported it from 
north of the 69" parallel, but this represents a large extension 
of its range southwards by some 7° of latitude. 


Carex lapponica O. F. Lang — YUKON. Sheldon Lake, 
131°13'W., 61°38'N. (UAC 70271). 

Cody et al. (1988, page 301) had earlier collected it in the 
extreme southeast of the Territory. 


Carex magellanica Lamark ssp. irrigua (Wahlenburg) 
Hiitonen — YUKON. Watson Lake at 128°41'W., 60°03'N. 
(UAC 73050). 

It was previously collected further west and north (Cody, 
1996, page 157) and further east (Cody et al. 1998, page 302). 


Carex marina Dewey — YUKON. Minto Landing, at 
136°53'W., 62°03'N. (UAC 66965), as a rare compo- 
nent of the grassy open areas in the boreal forest along 
the river bank, close to the steep, grassy, south-facing 
slopes of the valley walls. 

This is presumably an extension of its limited range in 
adjacent south-central Alaska in the rain shadow of the 
Wrangel Mountains (Hultén, 1968, page 239). Cody (1996, 
page 157) only reported it from Herschel Island, the Lower 
Blow River delta (Cody et al. 1998, page 303), and the Mal- 
colm River delta. 


Carex pachystachya Chamisso ex Steudel — YUKON. 
Macmillan Pass between 1097 m and 1250 m at 
130°30'W., 63°21'N. and 130°06'W., 63°23'N. (UAC 
61619, 61620), Tombstone Mountain Campground, 
km 75, Dempster Highway, about 138°14'W., 64°30'N. 
(UAC 61615), and Dragon Lake, North Canol Road, 
131°20'W., 62°33'N. (UAC 70287). 


2004 


These sites represent a large range extension from the 


south Yukon (Cody 1996, page 162) to the northern limit of 


the Boreal Forest. Harris (1998, page 269) previously listed 
it as being present at km 161.7, Robert Campbell Highway. 
It’s rarity needs to be verified. 


JUNCACEAE 
Juncus stygius Linnaeus ssp. americanus (Buch.) 
Hultn — YUKON. Thermokarst Mounds, South Fork, 
Blackstone River, Dempster Highway at 138°22'W., 
64°48'N., July 1987 (UAC 70257). 

Cody (1996, page 190) previously reported it from the 
Keno Hill area, while Cody et al. (1998, page 305) found it 
at Coal River in the southeast Yukon. 


Literature Cited 

Cody, W. J. 1994. The flora of the Yukon Territory: Additions, 
range extensions and comments. Canadian Field-Natural- 
ist 108: 428-476. 

Cody, W. J. 1996. Flora of the Yukon Territory. NRC 
Research Press, Ottawa, Ontario, Canada. 643 pages. 

Cody, W. J. 2000. Flora of the Yukon Territory. 2" Edition, 
NRC Research Press, Ottawa, Ontario, Canada. 669 pages. 

Cody, W. J., C. E. Kennedy, and B. Bennett. 1998. New 
records of vascular plants in the Yukon Territory. Canadian 
Field-Naturalist 112: 289-328. 

Cody, W. J., C. E. Kennedy, and B. Bennett. 2000. New 
records of vascular plants in the Territory II. Canadian 
Field-Naturalist 114: 417-443. 

Cody, W. J., C. E. Kennedy, and B. Bennett. 2001. New 
records of vascular plants in the Yukon Territory III. 
Canadian Field-Naturalist 115: 301-322. 


NOTES 


267 


Cody, W. J., C. E. Kennedy, B. Bennett, and V. Loewan. 
2002. New records of vascular plants in the Yukon Terri- 
tory [V. Canadian Field-Naturalist 116: 446-474. 

Cody, W. J., C. E. Kennedy, B. Bennett, and J. Staniforth. 
2003. New records of vascular plants in the Yukon Terri- 
tory [V. Canadian Field-Naturalist 117: 278-301. 

Douglas, G. W., G. W. Argus, H. L. Dickson, and D. F. 
Brunton. 1981. The rare vascular plants of the Yukon. 
Syllogeus #28. National Museums of Canada, Ottawa. 
61 pages. 

Flora of North America Editorial Commitee. 2002. Flora 
of North America 23. Oxford University Press, New York. 
608 pages. 

Harris, S. A. 1998. Effects of vegetation cover on soil heat 
flux in the southern Yukon Territory. Erdkunde 52: 265-285. 

Harris, S. A. 2004. Source areas of North Cordilleran en- 
demic flora: Evidence from Sheep and Outpost Mountains, 
Kluane National Park, Yukon Territory. Erdkunde 58: 
62-81. 

Hultén, E. 1968. Flora of Alaska and neighbouring Terri- 
tories. Stanford University Press, Stanford. 1008 pages. 
Moss, E. H. 1983. Flora of Alberta. 24 Edition, revised by 
John G. Packer. University of Toronto Press, Toronto. 687 

pages. 

Wahl, H. E., D. B. Fraser, A. C. Harvey, and J. B. Maxwell. 
1987. Climate of Yukon. Climatological Studies #40. 
Atmospherric Environment Service, Environment Canada. 
323 pages. 


Received 21 February 2002 
Accepted 21 August 2004 


“Ashkui’ Vernal Ice-cover Phenomena and Their Ecological Role in 


Southern Labrador 


SHAUNA M. BAILLie!, CORINNE D. WILKERSON? and TINA L. NEWBURY? 


‘Biology Department, P.O. Box 5000, St. Francis Xavier University, Antigonish, Nova Scotia B2G 2W5; e-mail:: sbaillie@ 
stfx.ca 

* Biology Department, Memorial University of Newfoundland, St. John’s, Newfoundland A1B 3X9; e-mail: r34cdw @mun.ca 

3 10 Highland Avenue, Corner Brook, Newfoundland A2H 2Y5; e-mail: tinalnewbury @hotmail.com 


Baillie, Shauna M., Corinne D. Wilkerson, and L. Tina Newbury. 2004. “Ashkui’” vernal ice-cover phenomena and their ecological 
role in southern Labrador 2002. Canadian Field-Naturalist 118(2): 267-269. 


This is the first documented incident of River Otter (Lutra canadensis) feeding on Common Goldeneye (Bucephala clangula) 
in a little studied region, southern Labrador. Our observations were made during spring staging when waterfowl aggregate 
at open water sites in frozen lakes and rivers, locally known as ashkui. We suggest that otters and raptors opportunistically 
forage on staging waterfowl at ashkui. 


Key Words: River Otter, Lutra canadensis, Common Goldeneye, Bucephala, clangula, Bald Eagle, Haliaeetus leuco- 


cephalus, predator-prey interactions, staging waterfowl, ashkui, Labrador. 


Ashkui (singular and plural form) is the Innu name 
given to sites of open water in river and lake systems 
within the frozen spring landscape of Labrador (Flet- 
cher and Breeze 2000*). Migratory waterfowl, includ- 
ing Common Goldeneye (Bucephala clangula), use 
ashkui as staging areas enroute to their breeding 
grounds. These birds arrive in groups of tens to hun- 
dreds to rest and rebuild energy reserves by feeding on 


invertebrates, fish, seeds and other plant material at 
ashkui (Newbury 2002*). Beaver (Castor canadensis), 
River Otter (Lutra canadensis) and Muskrat (Ondatra 
zibethicus), have been observed at these sites. Osprey 
(Pandion haliaetus) and Bald Eagle (Haliaeetus leu- 
cocephalus) are known to fish at ashkui (Fletcher and 
Breeze 2000*). The temporal existence, number and 
distribution of ashkui sites are influenced by a high 


268 


degree of inter-annual variability in the onset of spring 
thaw. As the winter snows disintegrate and waterfowl 
await availability of breeding habitat, they may be sus- 
ceptible to predation. In 2002, the opening of ashkui 
occurred two weeks later than in previous years (un- 
published data, Environment Canada, St. John’s, New- 
foundland). 

Our observations were made during a study exam- 
ining the effects of military low level flying on the 
behaviour of staging waterfowl at Lac Fourmont in 
southern Labrador (52°00'N, 60°15'W; Newbury 
2004*). While conducting waterfowl behavioural ob- 
servations, we witnessed a River Otter feeding on a 
male Common Goldeneye. We also observed a Bald 
Eagle attack another goldeneye resting on the ice. 


Study Area 

Lac Fourmont is situated within the Boreal Forest 
Region of eastern Canada and is influenced by con- 
tinental climatic regimes (Lopoukhine et al. 1977*). 
This region is covered with ice and snow until daily 
spring thaw temperatures rise to 5.1°C in May (30 year 
average for Goose Bay, Labrador; Environment Canada 
Meteorological Service, St. John’s, Newfoundland). 
Daytime average temperatures were 4.1°C in May 
2002 and ranged from 4.0 to 9.0°C for May during 
the years 1993 to 2002. River width at the observa- 
tion location was approximately | km, and the ashkui 
site was 20 m in breadth. Ice breadth from ashkui to 
shoreline was approximately 50 m. Forest stands were 
composed of mainly Black Spruce (Picea mariana) 
and Balsam Fir (Abies balsamea) interspersed with 
approximately 10 % Paper Birch (Betula papyrifera), 
Balsam Poplar (Populus balsamifera), and Trembling 
Aspen (Populus tremuloides). Lake-edge shrubs con- 
sisted of willows (Salix spp.) and alders (Alnus rugosa 
and A. crispa). River and bank substrate was composed 
of coarse-grained sand and pebbles with exposed boul- 
ders and bedrock of gneiss and mafic intrusions. River 
basin and landscape were low gradient with shallow 
sloping banks, wide flood plain and low relief hills. 

We conducted daily observations on waterfowl be- 
haviour from blinds using a Swarovski 60x spotting 
scope from 9:00 to 12:00 and 13:00 to 16:00 between 
26 April and 27 May 2002. Also, we conducted thir- 
teen random 1.5 h watches during dawn and dusk. 


Observations 

Fourteen species of waterfowl occupied the Lac 
Fourmont ashkui. Daily waterfowl numbers at the 
ashkui ranged from 34 to 376 during this study. A 
single River Otter was noted at the ashkui during at 
least five observation days. The number of sightings of 
mature and immature Bald Eagles, Osprey and Rough- 
legged Hawk (Buteo lagopus) totaled 7, 2, 1 and 1, 
respectively, over 8 non-consecutive study days. 

On 6 May 2002 at 11:35, we observed a River Otter 
grasping a male Common Goldeneye on the ice, ap- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


proximately 70 m from our observation blind on the 
south side of the ashkui. Although the capture was not 
observed, the goldeneye was alive and struggling when 
the otter was initially observed. The otter sat on its 
rear haunches with the goldeneye braced in its fore- 
paws and consumed the head of the duck first before 
eating the breast muscles and abdominal viscera. After 
approximately 10 minutes of feeding, the otter slipped 
into the water and swam downstream without the car- 
cass remains. We did not observe the fate of the duck 
carcass, as it was no longer visible on the ice when 
the otter left the feeding site. The bird flock, during 
this observation period, comprised mainly 65 Com- 
mon Goldeneyes, 27 Black Ducks, 113 Canada Geese 
(Branta canadensis). 

On 18 May 2002 at 16:03, we observed a Bald 
Eagle circle the ashkui and plunge downward toward 
a small group of Common Goldeneyes resting and 
sleeping on the ice, approximately 150 m from our 
position. Upon impact, the eagle initially captured a 
goldeneye. The eagle then struggled with the golden- 
eye while standing on the ice edge but was unable to 
maintain its grasp. The goldeneye fell from the ice to 
the water then became inter-mixed with the flock. This 
interaction lasted approximately one minute and, we 
were unable to identify the attacked goldeneye after 
its escape. The eagle then flew to the other side of the 
ashkui and rested on the ice. The bird flock, during 
this observation, comprised mainly 7 Common Gol- 
deneyes, 29 Black Ducks (Anas rubripes), 67 Canada 
Geese (Branta canadensis), 11 Greater Scaup (Aythya 
marila) and 20 Ring-necked Ducks (Aythya collards). 


Discussion 

Fish are often the main prey of River Otter and rap- 
torial predators (Chubbs and Trimper 1998; Lariviére 
and Walton 1998; Fletcher and Breeze 2000*; Heath 
et al. 2001; Jedrzejewska et al. 2001). Studies in the 
Great Lakes and North American boreal ecosystems 
have shown that the diet of River Otter comprises 
mostly fish (Lariviere and Walton 1998). However, 
otters have been known to feed opportunistically on 
small mammals, molluscs, reptiles, birds and fruits 
(Lariviére and Walton 1998). In west-central Idaho, 
otters supplemented their fish diet with invertebrates 
and reptiles (Melquist and Hornocker 1983). In Great 
Britain and Ireland, otters consume primarily fish, and 
lesser amounts of small mammals, medium-sized mam- 
mals, birds, herpetofauna, earthworms, other inverte- 
brates and carrion of large mammals (McDonald 2002). 
Jedrzejewska et al. (2001) showed that otters special- 
ize on prey taken from water. It is plausible that an 
otter fishing for fish under water may opportunisti- 
cally capture diving goldeneyes. Although River Otters 
were known to occasionally feed on birds (Lariviére 
and Walton 1998), we believe that this was the first 
documented case of an otter feeding on a Common 
Goldeneye. 


2004 


Instances of Bald Eagles preying on duck species 
are not rare (Jackman et al. 1999; Heath et al. 2001). 


_ Although fish dominate their diet, Bald Eagles have 


been reported to modify foraging behaviour and may 
forage opportunistically on staging waterfowl in situa- 
tions where return on such effort makes them econom- 
ical (Brown et al. 1998; Jackman et al. 1999). 

The temporal existence, number and distribution of 
ashkui, in southern Labrador during spring, may influ- 
ence the spatial and temporal distribution of piscivo- 
rous predators and migratory waterfowl by limiting 
the availability of open-water feeding and resting areas. 
Migratory waterfowl arriving from southern destina- 
tions must accumulate in greater concentrations when 
ashkui openings are limited. Though the behaviour 
and distribution of River Otter were not documented 
for Labrador, Lariviére and Walton (1998) stated that 
otters made heavy use of ashkui in winter and “almost 
exclusively” used open water for locomotion and 


_ foraging (Madsen and Prang 2001; Ruiz-Olmo et al. 


2001). Piscivorous raptors returning to breeding areas 
must travel to and congregate at available ashkui to 


_ fish. It was likely that spatial and temporal distribution 


of ashkui also affected predator-prey interactions. 

Our observations are examples of opportunistic for- 
aging events on waterfowl by predators that appear to 
use ashkui regardless of the presence of waterfowl. 
Further investigations in Labrador are needed to under- 
stand better the ecological importance of the tem- 
poral and spatial distribution of ashkui to mammalian 
and raptorial predators. 


Acknowledgments 

This project was funded in partnership by the In- 
stitute for Environmental Monitoring and Research 
(IEMR), the Canadian Wildlife Service (CWS), McGill 
University and the Department of National Defence 
(DND), and by the Northern Scientific Training Pro- 
gram. Many thanks to Sean Sharpe (IEMR), Tony 
Chubbs (DND) and Roger Titman (McGill University) 
for invaluable discussion, advice and/or informal manu- 
script review. Jean Sealy (CWS), Scott Gilliland (CWS) 
and Dawn Laing (DND) helped us with literature 
searches. Thanks to Keith Chaulk (CWS), Bruce Whif- 
fen (Environment Canada) and Sarah Hall (CWS) for 
providing information on weather and ice conditions. 


Documents Cited [marked * in text] 
Fletcher, C., and H. Breeze. 2000. Ashkui sites in the low- 
level flight training area, Labrador. Report prepared under 


NOTES 


269 


contract to Institute for Environmental Monitoring and 
Research (1.E.M.R.). Gorsebrook Research Institute for 
Atlantic Canada Studies, Saint Mary’s University, Halifax, 
Nova Scotia. 22 pages. 

Lopoukhine, N., N. A. Prout, and H. E. Hirvonen. 1977. 
The ecological land classification of Labrador. Ecologi- 
cal Land Classification Series, Number 4. Fisheries and 
Environment Canada. Halifax, Nova Scotia. 

Newbury, Tina L. 2002. Effect of low-level flights on the 
behaviour of spring staging waterfowl in Central Labrador. 
M.Sc. thesis Proposal. McGill University. Ste. Anne-de- 
Bellevue, Quebec, Canada. 

Newbury, Tina L. 2004. Effect of low-level military jet over- 
flights on the behaviour of spring-staging waterfowl at Lac 
Fourmont ashkui, Labrador, Canada. M.Sc. thesis. McGill 
University. Ste. Anne-de-Bellevue, Quebec, Canada. 


Literature Cited 

Brown, B. T., L. E. Stevens, and T. A. Yates. 1998. Influ- 
ences of fluctuating riverflows on Bald Eagle foraging 
behaviour. Condor 100: 745-748. 

Chubbs, Tony E., and Perry G. Trimper. 1998 The diet of 
nesting Ospreys, Pandion haliaetus, in Labrador. Canadian 
Field-Naturalist 112: 502-505. 

Heath, Joel P., Geoff Goodyear, and Joe Brazil. 2001. 
Observation of a Golden Eagle, Aquila chrysaetos, attack 
on a Harlequin Duck, Histrionicus histrionicus, in northern 
Labrador. Canadian Field-Naturalist 115: 515-516. 

Jackman, R. E., W. G. Hunt, J. M. Jenkins and P. J. 
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California. Journal of Raptor Research 332: 87-96. 

Jedrzejewska B., V. E. Sidorovich, M. M. Pikulik, and W. 
Jedrejewski. 2001. Feeding habits of the otter’and the 
American mink in Bialowieza Primeval Forest (Poland) 
compared to other Eurasian populations. Ecography 242: 
165-180. 

Lariviere. Serge, and Lyle R. Walton. 1998. Lutra cana- 
densis. Mammalian Species (587) American Society of 
Mammalogists. Pages 1-8. 

Madsen, A. B., and A. Prang. 2001 Habitat factors and the 

presence or absence of Otters Lutra lutra in Denmark. Acta 

Theriologica 46: 171-179. 

McDonald, R. A. 2002. Resource partitioning among British 

and Irish mustelids. Journal of Animal Ecology 71: 185- 

200. 

Melquist, W. E. and M. G. Hornocker. 1983. Ecology of 
river otters in west central Idaho. Wildlife Monographs 83: 
1-60. 

Ruiz-Almo. J., J. M. Lopez-Martin, and S. Palazon. 2001. 
The influence of fish abundance on the otter (Lutra lutra) 
population in Iberian Mediterranean habitats. Journal of 
Zoology 254: 325-336. 


Received 28 June 2002 
Accepted 19 October 2004 


270 THE CANADIAN FIELD-NATURALIST Vol. 118 


Sea Otter, Enhydra lutris, Sightings off Haida Gwaii / Queen 
Charlotte Islands, British Columbia, 1972-2002 


KIMBERLY RAUM-SURYAN!3, KENNETH PITCHER!, and RICHARD LAMY? 


' Alaska Department of Fish and Game, Division of Wildlife Conservation, 525 W. 67 Avenue, Anchorage, Alaska 99518 USA; 
e-mail: kraumsuryan @charter.net 

? Gwaii Haanas National Park Reserve/Haida Heritage Site, P.O. Box 37, 120 Second Avenue, Queen Charlotte, British Columbia 
VOT 1SO Canada 

>Current address: 928 NW Cottage Street, Newport, Oregon 97365 USA 


Raum-Suryan, Kimberly, Kenneth Pitcher, and Richard Lamy. 2004. Sea Otter, Enhydra lutris, sightings off Haida Gwaii/Queen 
Charlotte Islands, British Columbia, 1972-2002. Canadian Field-Naturalist 118(2): 270-272. 


On 27 June 2001 we observed and photographed a Sea Otter (Enhydra lutris) adjacent to a Steller Sea Lion (Eumetopias 
jubatus) haulout near Sgang Gwaay (Anthony Island), Haida Gwaii / Queen Charlotte Islands. This is one of only eight 
documented sightings of Sea Otters in these waters during the past 30 years. These sightings may represent the beginning of 


the expansion of Sea Otters to their former range off Haida Gwaii. 


Key Words: Sea Otter, Enhydra lutris, Haida Gwaii, Queen Charlotte Islands, British Columbia. 


The Sea Otter (Enhydra lutris) once ranged in near- 
shore waters along the North Pacific Rim from Japan 
to Baja California. The worldwide population of Sea 
Otters in the early 1700s was estimated to be between 
150 000 (Kenyon 1969) and 300 000 animals (John- 
son 1982). Middens (human food waste mounds) in- 
dicate that aboriginal people, including the Haida, 
hunted Sea Otters (Simenstad et al. 1978; Acheson 
1998; Sloan 2004). Although this harvest may have 
caused local reductions of Sea Otter populations near 
village sites (Simenstad et al. 1978), the species was 
abundant throughout its range prior to commercial ex- 
ploitation. Following the advent of commercial hunting 
in 1741, the worldwide population of Sea Otters de- 
clined to less than 2000 animals by 1911, limited to 
remnant groups throughout the range (Kenyon 1969). 
Many of these remnant groups (including one group 
located off the Haida Gwaii / Queen Charlotte Islands, 
British Columbia, Canada) were extirpated, likely due 
to their small size (Watson et al. 1997). The last Sea 
Otter recorded from Haida Gwaii was before 1920 
(Kenyon 1969) and the last recorded in British Colum- 
bia, near Vancouver Island (prior to reintroduction), 
was in 1929 (Cowan and Guiguet 1960; Kenyon 1969). 

In 1911 an International Treaty provided protection 
of Sea Otters from additional exploitation. Furthermore, 
to aid in the recovery of Sea Otters, various translo- 
cation projects were conducted from the 1950s to 
early 1970s in the Pacific coastal United States and 
Canada (Kenyon and Spencer 1960; Kenyon 1969; 
Burris and McKnight 1973; Bigg and MacAskie 1978; 
Jameson et al. 1982; Jameson et al. 1986; Riedman 
and Estes 1990). Since reintroduction, the Sea Otter 
population in British Columbia has increased 18.6% 
per year from 70 animals in 1977 to an estimated 2 500 
animals in 1998 (Watson 2000). Although Sea Otters 
continue to expand their range along Vancouver Island 
and the central coast of British Columbia (Watson 


2000; Figure 1), sightings off Haida Gwaii / Queen 
Charlotte Islands remain rare. 

On 27 June 2001 we observed and photographed a 
Sea Otter of unknown sex adjacent to a Steller Sea Lion 
haulout (52°04.966’ N, 131°13.811° W) just south of 
Sgang Gwaay (Anthony Island), Haida Gwaii / Queen 
Charlotte Islands (Figure 1). The Sea Otter was swim- 
ming on its back eating a sea urchin (Strongylocentrotus 
sp.). We observed the otter for approximately 30 min- 
utes as it swam and dove. This observation is one of 
only eight documented sightings (Edie 1973; Taylor 
and Gough 1977; Heise et al. 2003) of Sea Otters in 
waters surrounding Haida Gwaii / Queen Charlotte 
Islands during the past 30 years (Figure 1). 

This sighting raises some interesting questions 
regarding the origin of this individual and whether or 
not Sea Otters are expanding their range to include 
the Haida Gwaii / Queen Charlotte Islands once again. 
Kenyon (1969) suggested that the most significant 
factor limiting the spread of Sea Otter populations is a 
tendency of individual Sea Otters to occupy a limited 
home range. Although Sea Otters are not known to 
migrate, as populations increase, males generally are 
the first to explore new areas and their presence some- 
times indicates imminent range reoccupation (Loughlin 
1980; Garshelis et al. 1984; Pitcher 1989). Garshelis 
and Garshelis (1984) recorded long-distance move- 
ments of > 100 km for five adult territorial males. 
They postulated that the Sea Otters might have left 
an area of limited food resources to travel to an area 
where food was more abundant. When Sea Otters were 
extirpated, many of their primary benthic invertebrate 
prey became larger and more abundant (Estes and 
Palmisano 1974; Garshelis et al. 1986). When Sea Otters 
recolonize they initially encounter more abundant and 
larger prey resources than may be expected to persist, 
leading to densities of Sea Otters in recovering popu- 
lations greater than those that can be sustainable over 


Southeast 
Alaska 


NOTES 27) 


51 introduced in 1965 


International 
Boundary 


Haida Gwaii / 
Queen Charlotte 
Islands 


July 1999 


August 1990 
August 1976 
June 2001 
July 1972 


@ Sea Otter sightings 


A Sea Otter translocation sites 


C»> Current Sea Otter range 


@ August 2002 


55 introduced in 1965 


Ga British Columbia 


Wd 
D : 2 IN s a 


Q 
VY 


Goose Islands 
Group 


89 introduced between 1969 and 197 


200 Kilometers 


134° 132° 130° 128° 126° 


FicureE 1. Sightings of Sea Otters around Haida Gwaii / Queen Charlotte Islands, locations of Sea Otter translocations 


nearest Haida Gwaii, and current Sea Otter distribution in Southeast Alaska and British Columbia (map adapted 
from Parks Canada Report 038, Living marine legacy of Gwaii Haanas. IV: Marine mammal baseline to 2003 and 
marine mammal-related management issues throughout the Haida Gwaii region by Heise et al. 2003). 


time (Bodkin et al. 2000). As prey is consumed, Sea 
Otter densities should decline and fluctuate at an equil- 
ibrium density (Estes 1990) through decreased repro- 
duction, increased mortality, or through emigration 
(Bodkin et al. 2000). 

Sea Otter population increases in the Aleutian Islands 
were achieved largely by range expansion over deep, 


wide ocean passes (Estes 1990). Unless the Sea Otter 
we observed is a member of a previously undetected 
group off Haida Gwaii / Queen Charlotte Islands, this 
individual would have had to travel at least 175 km 
across an open expanse of water from the nearest 
known group of otters along the British Columbia coast 
(Figure 1). If this individual traveled from southeast 


272 


Alaska or northern Vancouver Island, the distance would 
be even greater. If Sea Otter populations have reached 
or exceeded carrying capacity in certain regions of 
British Columbia or southeast Alaska (Watson et al. 
1997; Watson 2000), this may be the beginning of the 
expansion of Sea Otters to their former range off Haida 
Gwaii / Queen Charlotte Islands. This range expansion 
may be particularly important given the precipitous 
population decline of Western Alaska Sea Otters dur- 
ing the past 15 years (Doroff et al. 2003) and the pro- 
posal to list the population as threatened under the 
United States Endangered Species Act (Federal Regis- 
ter: 11 February 2004; Volume 69, Number 28). 


Acknowledgments 

We thank the Gwaii Haanas National Park Reserve 
and Haida Heritage Site for allowing us to conduct 
Steller Sea Lion surveys (Research Permit 01-01) off 
Haida Gwaii, thereby allowing us to observe the Sea 
Otter mentioned. We are grateful to Jane Watson, Norm 
Sloan, Tom Gelatt, and two anonymous reviewers for 
their comments regarding this note. 


Literature Cited 

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People’ ]: a study of changing settlement strategies among 
the Kunghit Haida. British Archaelogical Reports Inter- 
national Series 711. 209 pages. 

Bigg, M. B., and I. B. MacAskie. 1978. Sea otters reestab- 
lished in British Columbia. Journal of Mammalogy 59: 
874-876. 

Bodkin, J. L., A. M. Burdin, and D. A. Ryazanov. 2000. 
Age- and sex-specific mortality and population structure 
in sea otters. Marine Mammal Science 16: 201-219. 

Burris, O. E., and D. E. McKnight. 1973. Game transplants 
in Alaska. Alaska Department of Fish and Game, Game 
Technical Bulletin (4). 

Cowan, I. M., and C. J. Guiguet. 1960. The mammals of 
British Columbia. British Columbia Provincial Museum 
Handbook (11). 

Doroff, A. M., J. E. Estes, M. T. Tinker, D. M. Burn, and 
T. J. Evans. 2003. Sea otter population declines in the 
Aleutian archipelago. Journal of Mammalogy 84: 55-64. 

Edie, A. G. 1973. Sea otter sighting at Cape St. James, Bri- 
tish Columbia. Syesis 6: 265. 

Estes, J. A. 1990. Growth and equilibrium in sea otter popu- 
lations. Journal of Animal Ecology 59: 385-401. 

Estes, J. A., and J. F. Palmisano. 1974. Sea otters: Their role 
in structuring nearshore communities. Science 185: 1058- 
1060. 

Garshelis, D. L., and J. A. Garshelis. 1984. Movements and 
management of sea otters in Alaska. Journal of Wildlife 
Management 48: 665-678. 

Garshelis, D. L., A. M. Johnson, and J. A. Garshelis. 1984. 
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Alaska. Canadian Journal of Zoology 62: 2648-2658. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Garshelis, D. L., J. A. Garshelis, and A. T. Kimker. 1986. 
Sea otter time budgets and prey relationships in Alaska. 
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Heise, K. A., N. A. Sloan, P. F. Olesiuk, P. M. Bartier, and 
J. K. B. Ford. 2003. Living marine legacy of Gwaii Haanas 
IV: Marine mammal baseline to 2003 and marine mammal- 
related management issues throughout the Haida Gwaii 
region. Parks Canada Technical Reports in Ecosystem 
Science Report 038. 150 pages. 

Jameson, R. J., K. W. Kenyon, A. M. Johnson, and H. M. 
Wight. 1982. History and status of translocated sea otter 
populations in North America. Wildlife Society Bulletin 
10: 100-107. 

Jameson, R. J., K. W. Kenyon, S. Jeffries, and G. R. 
VanBlaricom. 1986. Status of a translocated sea otter 
population and its habitat in Washington. The Murrelet 
67: 84-87. 

Johnson, A. M. 1982. Status of Alaska sea otter populations 
and developing conflicts with fisheries. Pages 293-299 in 
Proceedings of the Forty-seventh North America Wildlife 
Conference, Alaska. Wildlife Management Institute, Wash- 
ington, D.C. 

Kenyon, K. W., and D. L. Spencer. 1960. Sea otter popu- 
lation and transplant studies in Alaska, 1959. U.S. Fish 
and Wildlife Service Special Scientific Report — Wildlife 
49. 29 pages. 

Kenyon, K. W. 1969. The sea otter in the eastern Pacific 
Ocean. North American Fauna 68: 1-352. 

Loughlin, T. R. 1980. Home range and territoriality of sea 
otters near Monterey, California. Journal of Wildlife Man- 
agement 44: 576-582. 

Pitcher, K. W. 1989. Studies of the southern Alaska sea otter 
populations: Distribution, abundance, structure, range ex- 
pansion, and potential conflicts with shellfisheries. United 
States Fish and Wildlife Service Cooperative Agreement 
Number 14-16-009-954. 

Riedman, M. L., and J. A. Estes. 1990. The sea otter (En- 
hydra lutris): behavior, ecology and natural history. United 
States Fish and Wildlife Service Biological Report 90. 
126 pages. 

Simenstad, C. A., J. A. Estes, and K. W. Kenyon. 1978. 
Aleuts, sea otters, and alternate stable-state communities. 
Science 200: 403-411. 

Sloan, N. A. 2004. Northern abalone: Using an invertebrate 
to focus marine conservation ideas and values. Coastal 
Management 32: 129-143. 

Taylor, R., and B. Gough. 1977. New sighting of sea otter 
reported for Queen Charlotte Islands. Syesis 10: 177. 

Watson, J. C. 2000. The effects of sea otters (Enhydra lutris) 
on abalone (Haliotis spp.) populations. Pages 123-132 in 
Workshop on Rebuilding Abalone Stocks in British Col- 
umbia. Edited by A. Campbell. Canadian Special Publica- 
tion Fisheries and Aquatic Sciences 130. 

Watson, J. C., G. M. Ellis, T. G. Smith, and J. K. B. Ford. 
1997. Updated status of the sea otter, Enhydra lutris, in 
Canada. Canadian Field-Naturalist 111: 2: 277-286. 


Received 6 August 2002 
Accepted 26 August 2004 


ISABELLE SCHMELZER! and FRANK PHILLIPS” 


NOTES 


273 


_ First Record of a Barred Owl, Strix varia, in Labrador 


'Department of Environment and Conservation, Wildlife Division, Government of Newfoundland and Labrador. P.O. Box 2007, 
Corner Brook, Newfoundland A2H 7S1 Canada; e-mail: [sabelleSchmelzer@ gov.nl.ca 
"Department of Natural Resources, Forestry Division, Government of Newfoundland and Labrador, Goose Bay, Labrador 


AOP 1CO Canada 


Schmelzer, Isabelle, and Frank Phillips. 2004. First record of a Barred Owl, Strix varia, for Labrador. Canadian Field-Naturalist 


118(2): 273-276. 


A Barred Owl (Strix varia) was heard calling in central Labrador, Canada in 2001, and a dead owl was found in the same 
area in 2004. These are the first confirmed records for the Province of Newfoundland and Labrador. 


Key Words: Strix varia, Barred Owl, range, vagrant, Newfoundland and Labrador. 


Barred Owls (Strix varia) are common in southern 
Canadian forests from British Columbia through Nova 
Scotia (AOU 1998), occurring in mixed evergreen and 
mature deciduous woodlands, but also into the boreal 
forest in the northern portion of its range (Erskine 
1977; Godfrey 1986; Mazur et al. 1998). There are no 
confirmed records of Barred Owls for the Province of 
Newfoundland and Labrador (Todd 1963; ACCDC 
2004*). We document two occurrences of a Barred 
Owl near Happy Valley — Goose Bay, Labrador, three 
years apart. 

On the evening of 24 August 2001, at 22:00 hrs, a 
lone Barred Owl was heard calling in the forest north 
of Happy- Valley Goose Bay, Labrador, along the banks 
of the Goose River (53° 24'N, 60° 26'W) by one of 
the authors (IS) (Figure 1). The owl called continu- 
ously for several minutes from the same location, 
approximately 250 m away. The song was made up 
of the characteristic pattern of 8 hoots, separated into 
two parts by a pause after 4 hoots, and ending with a 
characteristic descending pitch on the last hoot (‘hoo- 
hoo hoohoo, hoohoo hoohooahhw’ ). The calls of this 
owl may be unfamiliar to residents of Labrador, but 
Barred Owl calls are very distinctive, and were imme- 
diately recognizable to one of the authors (IS). The 
only other owl species in the area with which the calls 
could possibly be confused is the Great Horned Owl 
(Bubo virginianus), whose call is of a deeper tone, and 
consists of five or six hoots, beginning with one em- 
phatic hoot, followed by several shorter and quieter 
calls, and ending with two long hoots on the same pitch 
(‘Hooo! hoohoohoo Hoooo, Hoooo’). The calls of the 
two owls differ in tone, pattern and rhythm, and are not 
difficult to tell apart by experienced listeners. 

Using a vocal imitation of the call, the author at- 
tempted to elicit a callback. The owl responded by 
leaving its perch, approaching her, and selecting a new 
perch within 50 m of her location. It made no other 
movements, and she was able to elicit vocal respons- 
es to her calls for nearly an hour. During this period, 
the owl occasionally varied its call either by repeat- 
ing only the last section ‘hoohooahhw’, or by short- 
ening the first section to a rising series of 3 notes 


followed immediately by the distinctive descending 
cackle (‘hoohoohoo hoohooahhw’ ). Unfortunately, due 
to darkness and heavy forest cover, the owl was not 
sighted. At the time of observation, the air temperature 
was 10°C, winds were calm, and the skies were clear. 
Tourists camped at the site reported hearing the same 
calls during the previous evening. 

On the evenings of 26, 28, and 31 August, at approx- 
imately 22:00, IS returned to the area to perform a 
nocturnal callback survey (using a vocal imitation of 
the call) to check for the presence of the owl. Roads 
and paths in forested areas within 10 km of the origi- 
nal sighting were surveyed, with stops every 0.3 — 
0.5 km, and several minutes spent at each’ station 
calling and listening for a response. No Barred owls 
were detected during these subsequent surveys. 

The second observation occurred three years later, 
in virtually the same location (53° 27'N 60° 18'W, on 
16 July 2004 (Figure 1). A dead owl was seen on the 
side of Route 530, at approximately 08:00 hrs. On 
closer inspection, the owl carcass showed no obvious 
sign of injury or illness, and was completely intact, 
with no sign of scavenging, and no fly larvae. The 
eyes were present and moist (not sunken, retracted, 
or absent as in older kills). Given the abundance of 
foxes (Vulpes vulpes), Ravens (Corvus corax), Gray 
Jays (Perisoreus canadensis) and other scavengers, it 
is the opinion of the authors that the owl had been 
dead for a few hours at most. 

Field marks on the owl included dark eyes, vertical 
barring on the belly and short, horizontal bars across 
the chest, all characteristic of Barred Owls (Figure 2). 
The owl was also assessed using the criteria of Pyle 
(1997). The lack of down on the head and contour 
feathers, flight feathers that were broad and uniform in 
color and wear, and the presence of squared bars on 
the primary coverts all suggest the bird was an adult 
(After Hatch Year or AHY). On dissection, the owl 
was determined to be a female by the presence of an 
ovary. No brood patch was present. 

Barred owls have been observed in Québec north 
of the St. Lawrence River (Gagnon and Bombardier 
1996). The breeding range has been reported to extend 


274 


Quebec 


Grebe 1992 


0) 150 300 450 


Kilometers Ve 
ay 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


FiGure 1. A summary of Barred Owl records in Labrador and nearby Quebec. Further description of the records is given in 


the text. 


to 50° North in eastern Canada (David 1980; Godfrey 
1986). Our review of the literature, however, revealed 
several sightings north of this latitude in Québec. For 
example, a Barred Owl was reported near Lac aux 
Cédres, 52° 00'N, 67° 07'W, near the Labrador border 
in 1917 (Todd 1963). It has been reported more recently 
further west near Nemiscau, 51° 42'N, 76° 15'W, and 
Lake Evans, 50° 55'N, 77° 00'W, (Grebe 1992). Addi- 
tionally, Comeau referred to the bird as “tolerably 
common” near Godbout, 49° 19'N, 67° 18'W, on the 
North Shore of the Gulf of St. Lawrence (in Merriam 
1882), and a specimen was captured at Lac Paterson 
in Roberval County (50° 11'N, longitude not specified) 
in 1954 (Godfrey 1957). Nonetheless, the occurrence 
of the owl in central Labrador is 500 km distant from 
the nearest previous report, and 700 km outside regions 
where the species is noted regularly. 

During the past several decades, the geographic 
range of the Barred Owl has expanded in western 
North America, and most recent discussion has focused 
on its movements into the Pacific Northwest (Levy 
1999; Boxall and Stepney 1982; Wright and Hay- 
ward 1998). However, on the basis of sightings in 
Quebec north of the St. Lawrence River, Harper (1958) 


proposed that the range of the Barred Owl was also 
extending northward in eastern North America. Al- 
though the Gulf of St. Lawrence and the Strait of 
Belle Isle present significant barriers to occurrence 
on the island of Newfoundland, the forests of Quebec 
and Labrador are continuous and present no obsta- 
cles to movement. The environs of Happy Valley — 
Goose Bay, including the Churchill River valley and 
coastal plain surrounding Lake Melville, belong to 
the “High Boreal Forest’ ecoregions, (Government of 
Newfoundland and Labrador 2004*) and consist of 
mature, dense mixed deciduous/coniferous forest on 
sandy soil. Predominant tree species are Trembling 
Aspen (Populus tremuloides) and White Birch (Betula 
papyrifera), with coniferous species such as Balsam 
Fir (Abies balsamea), and White and Black spruce 
(Picea glauca, P. mariana) occurring secondarily. 
Suitable breeding habitat for Barred Owls is often 
associated with the presence of snags used as nest 
cavities (Haney 1997), but also nests of other raptors 
or corvids (Elderkin 1987). This region features the 
most favorable climate in all of Labrador, including a 
growing season of 120-140 days. Conversely, the up- 
land plateaus of central and western Labrador are 


2004 


FiGureE 2: Adult female Barred Owl collected near Happy 
Valley — Goose Bay in July 2004, which showed char- 
acteristic gray-brown coloration, yellow bill, vertical 
barring on the belly and horizontal barring on the chest. 
Flight feathers (right wing shown) are broad and uni- 
form in colour and wear, suggesting an adult bird 
that has not yet molted. 


characterized by a subarctic forest and climate, and 
border the former ecoregion to the north and west. 
The elevation, vegetative communities, and severe cli- 
mate may pose a barrier to an eastward expansion from 
northern Québec. To the south and east however, broad 
river valleys and mixedwood boreal forests predomi- 
nate. 

In the absence of evidence of breeding, it is un- 
known whether this record represents a possible range 
expansion by the Barred Owl, or simply documents 
the occurrence of a vagrant bird in potentially suit- 
able habitat on two separate occasions. The fact that 
in One instance the owl was an adult female, and that 
she was found within the breeding season, does not 


NOTES 


275 


preclude range expansion as a possible explanation. 
The absence of geographical barriers to movement 
from northeastern Quebec, particularly from the north 
shore of the St. Lawrence, into south-central Labrador 
supports this notion. The establishment of owl surveys 
in Labrador should help to clarify such questions, and 
others, in the future. 


Acknowledgements 

We extend thanks to John Thomas and Joe Brazil for 
sharing their knowledge of bird records for Labrador, 
and for encouraging us to contribute and document 
our own sightings. Bruce Rodrigues used his banding 
expertise to sex and age the owl specimen. Chris 
Earley, Paul Linegar, and Bruce Mactavish provided 
constructive and useful comments on earlier drafts of 
the manuscript. 


Documents Cited (marked * in text) 

Atlantic Canada Conservation Data Center. 2004. 
Species Rarity Ranks: Labrador Birds, August 2004. P.O. 
Box 6416, Sackville, New Brunswick. E4L 1G6 http:// 
www.accde. com. 

Government of Newfoundland and Labrador. 2004. 
Department of Natural Resources-Forestry. Ecoregions of 
Labrador. http://www.gov.nf.ca/forestry/maps/eco_lab.stm. 

GREBE. 1992. Compexe Grande-Baleine. Avant-projet, 
phase II. Compte réndu de I’ étude télémétrique de I’ Aigle 
royal réalisée dans le complexe Grande Baleine en juin 
1992; mission G4. Report for Hydro Québec vice prési- 
dence environnement Montréal. 63 pages. 


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Vol. 118 


A. Comeau. Bulletin of the Nuttall Ornithological Club 
8: 244-245. 

Pyle, P. 1997. Identification guide to North American birds 
— part 1. Slate Creek Press, Bolinas, California, USA. 

Todd, W. E. C. 1963. Birds of the Labrador Peninsula and 
Adjacent Areas. A distributional list. University of Toronto 
Press, Toronto. 819 pages. 

Wright, A. L., and G. D. Hayward. 1998. Barred owl range 
expansion into the central Idaho wilderness. Journal of 
Raptor Research 32: 77-81. 


Received 23 January 2002 
Accepted 30 August 2004 


Forked Three-awned Grass, Aristida basiramea Engelm. ex Vasey: 
A New Addition to the Flora of Quebec 


JACQUES BRISSON 


Institut de recherche en biologie végétale, Université de Montréal, 4101, Sherbrooke Street East, Montréal, Québec H1X 2B2 
Canada 


Brisson, Jacques. 2004. Forked Three-awned Grass, Aristida basiramea Engelm. ex Vasey: a new addition to the flora of Quebec. 
Canadian Field-Naturalist 118(2): 276-277. 


A population of Forked Three-awned Grass (Aristida basiramea Engelm. ex Vasey; Poaceae) was found for the first time in 
Quebec, on a sand barren of the Cazaville region (Haut-Saint-Laurent). The only other region where this species is known 
in Canada is on the southern side of Georgian Bay in Ontario. 


Key Words: Aristida basiramea, Forked Three-awned Grass, Poaceae, rare plants, Quebec 


Une population d’Aristida basiramea Engelm. ex Vasey (Poaceae) fut trouvée pour la premiére fois au Québec, dans une 
lande sableuse de la région de Cazaville (Haut-Saint-Laurent). Le seul autre endroit ot: l’espéce est présente au Canada est 
sur le coté sud de la Baie Georgienne en Ontario. 


Mots-clés : Aristida basiramea, Poaceae, plantes rares, Québec 


In September 2001, during an ecological survey near 
Cazaville, in the Haut-Saint-Laurent region of south- 
ern Quebec, a population of Forked Three-awned Grass 
(Aristida basiramea Engelm. ex Vasey) was found in 
a dry, sandy grass-field. Other populations were later 
located in similar habitats nearby. This is the first 
time the species has been reported for Quebec. In 
Canada, A. basiramea is known from four extant 
naturally-occurring sites located in southern Ontario, 
three of which are in Simcoe County, and one in 
Muskoka County (Allen 2001). The extent of 
occurrence in Ontario only totals 16 hectares. There 
is also one adventive station in northwestern Ontario, 
at Rainy River (Allen 2001). A few other reports 
exist, such as one in Norfolk County in Ontario, and a 
few more in Manitoba. However, the locality of the 
Norfolk specimen may be the result of a labelling 
error (Reznicek 1984), while the reports from 
Manitoba are considered questionable due to the 
absence of existing specimens (Scoggan 1957; Allen 
2001). Thus, the Cazaville area becomes the second 
area in Canada where the presence of the species is 
confirmed. The species is listed as rare in Ontario 
(Reznicek 1984) and is considered threatened in Can- 


ada (COSEWIC 2003). On the basis of the newly 
discovered population, A. basiramea should be added 
to the list of rare vascular plants likely to be designated 
threatened or vulnerable (Labrecque and Lavoie 2002). 

The genus Aristida is represented by 250 to 300 spe- 
cies, 29 of which are native to North America north 
of Mexico (Flora of North America Editorial Com- 
mittee 2003). A. basiramea is an annual plant from 
30 to 60 cm high characterized by glumes of unequal 
length and 1-flowered spikelets terminated by three 
long awns, one in the middle with a twisted base and 
two shorter straight awns on each side. It is abundant 
on dry sterile or sandy soil in the midwestern states. 
At the northeastern periphery of its range in the United 
States, it forms disjunct populations on dry lands and 
along sandy roadsides, some populations of which ap- 
pear to be adventive. The closest reports of A. basira- 
mea south of the border with Québec were from Platts- 
burg, Clinton Co., New York (S. J. Smith, 25 July 1965, 
NYS); Columbia, northern Coos Co., New Hampshire 
(A. S. Pease, 17 September 1955, NEBC); and Avon, 
Franklin Co., Maine (A. Haines, 26 October 1990, 
MAINE). The species is recognized as rare in Maine, 
Iowa and Colorado (Allen 2001). 


The Cazaville area (45°03’ N, 74°22’ W) is located 
in the Mixed Plain Ecozone of the St-Lawrence Low- 
land Ecoregion (Ecological Stratification Working 
Group 1995). The area is characterized by a vast sandy 
plain of littoral origin dating from the last post-gla- 
cial period. White Pine (Pinus strobus) forests were 
abundant before European settlement (Brisson and 
Bouchard 2003), but today, the area is occupied by 
sandy, grassy fields, sand barrens and secondary forests 
of Red Maple (Acer rubrum), Trembling Aspen (Popu- 
lus tremuloides) and Gray Birch (Betula populifolia). 
Aristida basiramea was found on open, sandy grass- 
field with Poa compressa, Danthonia spicata, and vari- 
ous types of lichens. The sandy plain of Cazaville is 
host to other rare plant species. The northernmost 
colony of Monarda punctata, a species rare for Quebec 
and Canada, was recently found nearby (Boudreault 
and Brisson 1994). As well, Hedeoma hispida, consid- 
ered rare for Quebec (Labrecque and Lavoie 2002), 
is also found on the sandy plain. 

Specimens of A. basiramea were deposited at the 
Marie-Victorin Herbarium (MT: Brisson Number JBO1- 
25) 


Acknowledgments 

I thank Stuart Hay (Marie-Victorin Herbarium) and 
Stephen Darbyshire (Agriculture and Agri-Food Can- 
ada), who reviewed the manuscript and provided useful 
information. I am also grateful to Richard Mitchell 
(Biological Survey, New York State Museum) and Ray 
Angelo (Harvard University Herbaria, New England 
Botanical Club) for kindly providing information on 
A. basiramea in the U.S. 


NOTES 


Documents Cited (marked * in text) 

Allen, G. M. 2001. COSEWIC Status Report on Forked 
Three-awned Grass. Committee on the Status of Endan- 
gered Wildlife in Canada. Environment Canada, Ottawa, 
Ontario. 26 pages. 


Literature Cited 

Boudreault, C., and J. Brisson. 1994. Une addition a la flore 
du Québec: Monarda punctata var. villicaulis (Lamiaceae). 
Canadian Field-Naturalist 108: 499-500. 

Brisson, J., and A. Bouchard. 2003. Human activities caused 
major changes in tree species composition in southern 
Quebec, Canada. Ecoscience 10: 236-246. 

COSEWIC. 2003. Canadian Species at Risk, November 
2003. Committee on the Status of Endangered Wildlife in 
Canada. 44 pages. 

Ecological Stratification Working Group. 1995. A National 
Ecological Framework for Canada. Agriculture and Agri- 
Food Canada, Research Branch, and Centre for Land and 
Biological Resources Research and Environment Canada, 
State of the Environment Directorate, Ecozone Analysis 
Branch, Ottawa/Hull. Report and National map at 1: 
7 500 000 scale. 

Flora of North America Editorial Committee. 2003. Flora 
of North America North of Mexico. Volume 25: Poaceae. 
(Part 2). New York and Oxford. 814 pages. 

Labrecque, J., and G. Lavoie. 2002. Les plantes vasculaires 
menacées ou vulnérables du Québec. Gouvernement du 
Québec, Ministére de l’environnement. Direction du patri- 
moine écologique et du développement durable, Québec, 
200 pages. 

Reznicek, A. A. 1984. Poaceae — Aristida basiramea . One 
page in Atlas of the rare vascular plants of Ontario. Edited 
by G. W. Argus and D. J. White. 1982-1987. National 
Museum of Natural Sciences. Looseleaf. 

Scoggan, H. J. 1957. Flora of Manitoba. National Museum 
of Canada, Bulletin 140. 619 pages. 


Received 30 November 2001 
Accepted 21 June 2004 


278 THE CANADIAN FIELD-NATURALIST Vol. 118 


Evidence for the Use of Vocalization to Coordinate the Killing of a 
White-Tailed Deer, Odocoileus virginianus, by Coyotes, Canis latrans 


ERICH M. Muntz! and BRENT R. PATTERSON? 


Department of Natural Resources, Wildlife Division, 136 Exhibition Street, Kentville, Nova Scotia B4N 4E5 Canada 

‘Current address: Parks Canada Agency, Cape Breton Highlands National Park of Canada, Ingonish Beach, Nova Scotia 
BOC 1L0 Canada 

*Current address: Ontario Ministry of Natural Resources, Wildlife Research and Development Section, 300 Water Street, 3" 
Floor North, Peterborough, Ontario K9J 8M5 Canada; e-mail: brent.patterson@mnr.gov.on.ca. Author to whom 
correspondence should be addressed. 


Muntz, Erich M., and Brent R. Patterson. 2004. Evidence for the use of vocalization to coordinate the killing of a White-Tailed 
deer, Odocoileus virginianus, by Coyotes, Canis latrans. Canadian Field-Naturalist 118(2): 278-280. 


Among the social canids, howling is largely accepted as playing a role in territory maintenance. However, its role in commu- 
nication within packs, such as announcing departures from den and rendezvous sites and coordinating reunions or movements, 
remains largely speculative. We report an observation where a radio-collared adult male Coyote (Canis latrans) and his mate 
seemed to summon two other Coyotes (presumed to be their offspring) from ~700 m away to join in the successful pursuit 
of an adult male White-tailed Deer (Odocoileus virginianus). Our observation suggests that Coyotes can use vocalization as 


an effective means of coordinating social activities such as the hunting of large prey. 


Key Words: Eastern Coyote, Canis latrans, predation, vocalization, social organization, Nova Scotia. 


Auditory communication serves an important role 
in the social ecology of the Canidae (Harrington and 
Mech 1978, 1979; Theberge and Falls 1967; Gese 
and Ruff 1998). Howling is a means of long-distance 
communication that apparently can be heard at dis- 
tances of > 6 km (McCarley 1975; Harrington and Mech 
1979). Among Coyotes (Canis latrans) and Wolves 
(Canis lupus), howling is largely accepted as playing 
a role in territory maintenance (Joslin 1967, Harrington 
and Mech 1978, 1979; Gese et al. 1988). Intrapack 
communicatory roles, such as announcing departures 
from den and rendezvous sites, and coordinating re- 
unions or movements remain largely speculative (The- 
berge and Falls 1967; Mech 1970; Harrington and 
Mech 1978). Bender et al. (1996) suggested that howl- 
ing among Coyotes might serve to reunify packs to 
facilitate the hunting of ungulates. Herein we report a 
case where a breeding pair of Coyotes in pursuit of a 
large White-tailed Deer (Odocoileus virginianus) was 
able to entice two other Coyotes to join the pursuit 
from ~700 m away. 


Observation 

We interpreted the details of the chase while snow- 
tracking a radio-collared Coyote and his mate on 27 
February 1994 as part of a study of the effects of the 
distribution and abundance of Snowshoe Hares, Lepus 
americanus, and White-tailed Deer on the life history 
of Coyotes in Nova Scotia (Patterson and Messier 
2000, 2001; Patterson et al. 1998). The actual event 
probably occurred during the previous night. 

Radio-collared Coyote AM3 and his mate were 
traveling west on a snow-covered (10-15 cm) secondary 
road | km east of Kejimkujik National Park (44°20'N, 


65°15'W) when they abruptly veered due north. 
Lengthened strides indicated that their pace increased, 
and after 40 m the pair jumped three deer, of which 
two headed west and the other east. Both Coyotes 
chased the deer that ran to the east. After 280 m the 
two Coyotes split up, with the larger radio-collared 
male chasing the deer down a steep bank into a bowl- 
shaped depression. AM3 then swung wide to the left 
of the deer, apparently in an attempt to steer the deer 
into the path of the other Coyote that had remained 
on the rim of the depression. The second Coyote cir- 
cled the depression and ran down the opposite side in 
front of the deer. We interpreted this as an effort to 
prevent the deer from leaving the depression. 

Within 3 m of the second Coyote’s resumption of 
the chase, both Coyotes attacked the deer and drew hair 
but no blood. The deer escaped and again attempted to 
run up the side of the depression when AM3 swung to 
the right and turned the deer back down into the depres- 
sion, where the second Coyote was waiting and again 
resumed the chase. The Coyotes attacked the deer again 
at 383 m where a larger area of snow was trampled 
down than during the first attack. There was more 
hair strewn about, but still no blood. At this point two 
more Coyotes became involved in the chase. At 390 m 
more hair was detected as well as blood, indicating 
another attack. The deer broke away once more only to 
be attacked again at 397 m. After escaping yet again 
the deer made a long run across the middle of the 
depression. Considerable amounts of blood and hair 
strewn about a 20 x 30 m packed-down area at 590 m 
indicated a more serious struggle. The deer made one 
final escape and was pulled down by the front end at 
630 m, with tracks indicating that the deer was drag- 


2004 


ging at least two of the Coyotes, which appear to have 
been hanging off its sides. After dragging the Coyotes 


~ for 40 m the deer was killed at the 670 m mark. 


Examination of the carcass revealed that the deer 
was a large 3.5 year old buck that showed no obvious 
debilitations and was apparently in good health (>80% 
femur marrow fat content and other visible body fat 
reserves). We believe that the deer would likely have 
escaped if it could have got out of the depression. The 
two Coyotes initially involved in the chase appeared 
to have trouble drawing blood from the deer until the 
other two Coyotes joined them. Backtracking later 
revealed that the two Coyotes that joined the chase in 
progress had been traveling in another direction when 
they abruptly turned and trotted 690 m in a direct line 
to join the chase. Forest cover was dense, precluding 
any possibility that these Coyotes observed the chase 
prior to joining in. AM3 and his mate were typically 
accompanied by two of their young of the year when 
traveling in winter 1994 (Patterson and Messier 2001). 
We believe that the two Coyotes that joined the chase 
“in progress” were probably these same juveniles that 
had been temporarily disassociated from their parents. 
We speculate that they must have heard AM3 or his 
mate howling or yipping and were able to determine 
that it would be to their benefit to join the breeding 
pair promptly. 


Discussion 

Although we can not verify that howling was used 
to draw the other two Coyotes to the scene of the chase, 
we can think of only one other means by which two 
Coyotes ~700 m away in forested cover may have been 
able to so directly and rapidly locate the scene of the 
chase. White-tailed Deer can snort loudly when alarmed, 
but snorts are generally only given when a deer per- 
ceives danger but does not feel directly threatened 
(Hirth and McCullough 1977; Marchinton and Hirth 
1984). Furthermore, snorts are more likely to be given 
by maternal family groups than by bucks (Hirth and 
McCullough 1977). Thus it seems unlikely that snort- 
ing by the buck alerted the other two Coyotes to the 
chase. 

Among forest-dwelling eastern Coyotes increased 
reproductive fitness and inclusive fitness for juveniles 
before dispersal seem to be the ultimate factors influ- 
encing group living (Messier and Barrette 1982; Pat- 
terson and Messier 2001). Increased efficiency at using 
large prey appears to be a secondary benefit (Gese et 
al. 1988; Messier and Barrette 1982; Patterson and 
Messier 2001). Coyotes in Nova Scotia were more suc- 
cessful at killing deer when thick snow cover impeded 
deer movements (Patterson and Messier 2000). There 
was only 10-15 cm of snow on the ground during the 
event described here and our observations suggest that 
in this particular incidence the snow cover may have 
been a hindrance to the Coyotes, thus benefiting the 


NOTES 


279 


deer. It was clear that AM3 and his mate were having 
difficulty subduing the deer on their own. We believe 
that they would not likely have been able to make phys- 
ical contact with the deer if it had not entered the 
depression. Although Patterson and Messier (2000) did 
not detect a consistent increase in deer killing rates 
for groups of 2-5 Coyotes, groups of >4 Coyotes 
killed proportionately more deer (Patterson 1999). We 
suspect that larger group sizes may be more advan- 
tageous in hunting large prey in the absence of other 
contributing factors such as thick snowcover or glare 
ice. The proximate mechanism is likely an increase 
in the probability of at least one member of the group 
making physical contact with the deer and slowing it 
down enough for other group members to assist in 
dispatching it. Our observation supports this idea and 
suggests that Coyotes can use howling as an effective 
means of coordinating social activities such as the 
hunting of large prey. 


Acknowledgments 

This work was supported by Parks Canada, the 
Nova Scotia Department of Natural Resources and 
Acadia University. We appreciate comments by D. O. 
Joly on an earlier draft of the manuscript. 


Literature Cited 

Bender, D. J., E. M. Bayne, and R. M. Brigham. 1996. Lunar 
condition influences coyote (Canis latrans) howling. 
American Midland Naturalist 136: 413-417. 

Gese, E. M., and R. L. Ruff. 1998. Howling by coyotes 
(Canis latrans): variation among social classes, seasons, 
and pack sizes. Canadian Journal of Zoology 76: 1037- 
1043. 

Gese, E. M., O. J. Rongstad, and W. R. Mytton. 1988. 
Relationship between coyote group size and diet in south- 
eastern Colorado. Journal of Wildlife Management 52: 
647-653. 

Harrington, F. H., and L. D. Mech. 1978. Howling at two 
Minnesota wolf pack summer homesites. Canadian Journal 
of Zoology 56: 2024-2028. 

Harrington, F. H., and L. D. Mech. 1979. Wolf howling and 
its role in territory maintenance. Behaviour 68: 207-249. 

Hirth, D. H., and D. R. McCullough. 1977. Evolution of 
alarm signals in ungulates with special reference to white- 
tailed deer. American Naturalist 111: 31-42. 

Joslin, P. W. B. 1967. Movements and homesites of timber 
wolves in Algonquin Park. American Zoologist 7: 279-288. 

Marchinton, R. L., and D. H. Hirth. 1984. Behavior. Pages 
129-168 in White-tailed deer: Ecology and Management. 
Edited by L. K. Halls. Stackpole Books, Harrisburg, 
Pennsylvania. 

McCarley, H. 1975. Long distance vocalizations of coyotes 
(Canis latrans). Journal of Mammalogy 56: 847-856. 
Mech, L. D. 1970. The wolf: the ecology and behavior of an 
endangered species. Natural History Press, New York. 

384 pages. 

Messier, F., and C. Barrette. 1982. The social system of the 
coyote (Canis latrans) in a forested habitat. Canadian Jour- 
nal of Zoology 60: 1743-1753. 


280 


Patterson, B. R. 1999. The effects of prey distribution and 
abundance on eastern coyote life history and predation 
on white-tailed deer. Ph.D. thesis, University of Saskat- 
chewan, Saskatoon, 204 pages. 

Patterson, B. R., L. K. Benjamin, and F. Messier. 1998. 
Prey Switching and feeding habits of Eastern Coyotes in 
Relation to Snowshoe Hare and White-Tailed Deer Densi- 
ties. Canadian Journal of Zoology 76: 1885-1897. 

Patterson, B. R., and F. Messier. 2000. Factors Influencing 
killing rates of White-Tailed Deer by Coyotes in Eastern 
Canada. Journal of Wildlife Management 64: 721-732. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Patterson, B. R., and F. Messier. 2001. Social organization 
and space use of Coyotes in Eastern Canada relative to 
prey distribution and abundance. Journal of Mammalogy 
82: 463-477. 

Theberge, J. B., and J. B. Falls. 1967. Howling as a means of 
communication in timber wolves. American Zoologist 7: 
331 -338. 


Received 24 September 2002 
Accepted 11 October 2004 


Book Reviews 


ZOOLOGY 


Fish of Alberta 


By Amanda Joynt and Michael G. Sullivan. 2003. Lone 
Pine Publishing, Edmonton, Alberta. 176 pages. $18.95. 
ISBN 1-55105-191-5. 


This book is a popular account of the 54 established 
and 11 more rare fishes of Alberta. It consists of 
Acknowledgements, Foreword (by J. S. Nelson, the 
expert on fishes of Alberta), Reference Guide, Intro- 
duction, Keys to the Fishes, Species Accounts, Size 
Comparisons, Other Alberta Fishes (the rarer species), 
Glossary, Checklist, Selected References, Further Infor- 
mation, Index of Scientific Names, Index of Com- 
mon Names and About the Authors. These are mostly 
self-explanatory. The Reference Guide is a colour 
pictorial summary of each of the 54 species while 
Size Comparisons show the fish in relation to a loonie 
or a page in the book. The Glossary, Selected Refer- 
ences and Further Information sections are short but 
give some additional sources and explications of terms. 
Schreckstoff is misspelled and the term “alarm sub- 
stance” is more user-friendly. Laterally compressed 
is a tautonym, as compressed means flattened from 
side to side. For a more complete listing of terms 
used in ichthyology see www.briancoad.com. 

The ichthyofauna includes 16 families with min- 
nows (Cyprinidae) having 17 species, 3 introduced; 
trouts (Salmonidae) 16 species, 6 introduced; and suck- 
ers (Catostomidae) 7 species. Other families are lam- 
preys, sturgeons, mooneyes, bullhead catfishes, pikes, 
trout-perches, cods, livebearers (2 species introduced 
to the Banff hot spring marshes), sticklebacks (1 
introduced), sculpins, sunfishes (introduced), perches 
and cichlids (1 introduced to the Banff marshes). 

The Introduction has information about identifying 
fishes, the lives of fishes, adaptations to life in water, 
the underwater world, drainage basins of Alberta, top 
Alberta fishwatching sites and fishwatching (“not yet 
a popular sport’), history of fishes and conservation 
issues. Key words are in bold text and some, but not 
all of these, have illustrations. One illustration, of Bull 
Trout and Walleye larvae used to explain reproductive 
strategies, is confusing in that these two distinctive 
species have the same larva illustrated. 

The Species Accounts comprise two pages for each 
established species, in landscape orientation. This 
requires some twisting for reading introductory and 
other sections that have a standard orientation. None- 
theless, the landscape arrangement enables the fish 
illustrations to spread across the broadest reach of 
the page. Each Account has an introductory section, 


Viewing Tips (localities and habitats where the fish 
can be seen), Feeding, Spawning, Other Names, Did 
You Know? (an anecdote or interesting factoid), ID 
(identification characters), Similar Species, Status, 
Habitat, Length and Weight, the colour illustration and 
a colour, shaded distribution map. 

Generally, the book is a good, popular treatment of 
this fish fauna written in a readable style that is not as 
dry as more scientific accounts. Some errors occur, per- 
haps as a result of this style or perhaps from the more 
pedantic viewpoint of the reviewer; e.g., in the Long- 
nose Sucker account pharyngeal teeth are said to be in 
the mouth which could mislead the reader into thinking 
these throat teeth are readily visible. 

The definition of “fish” and “fishes” is given on page 
10 in the Introduction, fish being a single individual 
or more than one individual of a single species while 
fishes refers to more than one species. Curiously, the 
book is wrongly titled. The Reference Guide has a col- 
our code for groups of species but since these colours 
are various shades of green and blue, they do not lend 
themselves to a ready means of locating species groups. 

The fish illustrations are positioned with the head to 
the right, not a problem for most users, but somewhat 
disconcerting to those familiar with most fish books 
where traditionally the head faces left. The colour illus- 
trations lack the details that can be included on line 
drawings, so certain critical features are not always 
visible. The illustrations have streaks of white meant as 
highlights but in some specimens look like a colour 
pattern. 

Scientific names are given for all species and are ac- 
curate and, although Stizostedion may now be correctly 
Sander, the decision to leave it as the more familiar 
North American genus is probably apposite for infor- 
mation retrieval. Conversely, the Arctic Lamprey is 
given its newer name, Lampetra camtschatica, which 
is less well known than the older version, L. japonica, 
which is not mentioned and would inhibit search for 
information in other books. Common names are stan- 
dard and include the Northern Pikeminnow, although 
its former name Northern Squawfish was not so much 
an “unpopular” designation as a derogatory one. 

Despite the minor criticisms outlined above, this 
book is a good introductory guide to Alberta’s fishes. 
Alberta seems well served in this respect with Nelson 
and Paetz’s earlier book (1992). The older work pro- 
vides a more detailed account of the fish fauna and 
may be more suited to the serious student of Alberta’s 


281 


fishes — there is an extensive list of references (24 
pages compared to only 11 references in Joynt and 
Sullivan). The book reviewed here has accounts for 
65 species while Nelson and Paetz have a more 
complete analysis of 59 species and a further 29 
species recorded as rare or doubtful for Alberta. 


Sharks 


By Andrea and Antonella Ferrari. 2002. Firefly Books, Tor- 
onto, Ontario. 256 pages. $24.95. ISBN 1-55209-629-7. 


Sharks are the most popular of fishes for publica- 
tions as people seem to have a fascination for organ- 
isms that can eat you (although more sharks are killed 
by people than the reverse). This is one of many that 
have appeared on this topic. 

Despite its title, the book covers the appearance and 
behaviour of 120 species of sharks and rays. Coverage 
is therefore not complete as sharks and rays number 
over 1000 species (as the Foreword states), the aim be- 
ing to give an overview of the main groups by select- 
ing typical and unusual or fascinating species. Four 
species of rabbitfishes (out of about 31 species in the 
Chimaeriformes) are also mentioned. 

The book has an Introduction of 77 pages with 
descriptions of anatomy and biology. Three pages are 
devoted, deservedly, to explaining the ampullae of 
Lorenzini, an important feature of shark anatomy and 
biology, first discovered by an Italian scientist in the 
seventeenth century and, surely by coincidence, this 
book has Italian authors and is a translation from an 
Italian version. This is followed by how to avoid sharks, 
how to assist them (by not eating them or buying shark 
teeth), marine organisms more dangerous than sharks 
(mostly venomous sea snakes, cone shells, sea urchins, 
jellyfishes and fishes, but also fish that bite divers en- 
thusiastically), and personal accounts of shark encoun- 
ters. The Entries section describes the sharks, rays and 
rabbitfishes (153 pages), the Appendices (17 pages) 
give a Classification of Sharks (but not rays or rabbit- 
fishes), an Index, Bibliography and Websites (each a 
page long), and Photographic Credits. The websites 
“Catalog of Fishes” and “FishBase” are not quoted, 
although these give entry to much of the names, biol- 
ogy and literature on sharks and rays world-wide for 
the more serious student. There are pictorial keys to 


In Search of the Golden Frog 


By Martha Crump. 2002. University of Chicago Press, 5801 
Ellis Avenue, Chicago, Illinois 60637 USA. xiv + 298 
pages, Cloth US$27. 


Any young person thinking of taking a degree in 
one of the biological sciences, particularly if they plan 
to become a field biologist, should read this book. 
The author, currently Adjunct Professor of Biology at 
Northern Arizona University, chronicles her life from 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Literature Cited 

Nelson, J. S., and M. J. Paetz. 1992. The Fishes of Alberta. The 
University of Alberta Press, Edmonton and the University of 
Calgary Press. 2"4 Edition, 437 pages. 


BRIAN W. COAD 


Canadian Museum of Nature, P.O. Box 3443, Station D, 
Ottawa, Ontario K1P 6P4 Canada 


the shark and ray orders but not any for families or 
species so the book cannot be used, nor is it meant, as 
a field guide. 

The Entries section describes each selected species, 
allotting half to two pages per species. There is an 
annoying symbol system at the top of each account 
to indicate when the shark is active and its danger 
level with respect to humans. This information could 
easily have been included in the text. A distribution 
map is given but at | x 2 cm can only convey a 
general impression of where these sharks and rays 
are found. The map for the Bluntnose Shark is in- 
accurate, for example, showing it in Canadian Arctic 
waters when it was first caught at its northern limit in 
Nova Scotian waters in 1989 (Gilhen and Coad 1991). 
The common and scientific names are given but the 
author and date of the scientific name is always in 
parentheses (here and in the Classification of Sharks). 
The authors (or text editor?) seem not to be aware 
that parentheses are only used when the species is 
placed in a genus other than the one it was originally 
described under. The text comprises the Family to 
which the shark or ray belongs, Range, Habitat, Size, 
and Habits. All parts of the book are richly illustrated 
with colour photographs, over 450 in all. 

Despite the comments above, this book is a good 
general introduction to sharks and rays. It is lavishly 
illustrated, as all such books must be, of a convenient 
size, and with a reasonable price. 


Literature Cited 

Gilhen, John, and Brian W. Coad. 1991. The bluntnose sixgill shark 
Hexanchus griseus (Bonnatere, 1788), new to the fish fauna of At- 
lantic Canada. Proceedings of the Nova Scotian Institute of Science, 
39 (1989): 75-77. 


BRIAN W. COAD 


Canadian Museum of Nature, P.O. Box 3443, Station D, 
Ottawa, Ontario K1P 6P4 Canada 


her postgraduate work in 1968 through various projects 
to 1998. The book concentrates on the field programs 
that are the real love of Professor Crump’s life. Her 
main interests are reptiles and amphibians, but her 
passion is frogs. 

Taking advantage of an opportunity offered by her 
discerning professor, she joins a team going to the 
Amazon section of Ecuador. From this exposure which 


2004 


reinforces her desire to work in the field, she returns 
| repeatedly to work in South America. Her narrative 
_ includes her excitement at being a scientist and mak- 
ing discoveries to contribute to her profession. She 
also chronicles some of the problems she encounters. 
These do not include frightening encounters with large 
or dangerous animals. The hazards fall into two cate- 
gories. The small pests, mosquitoes, chiggers, chigoes, 
bot flies and ants, causes plenty of discomfort. But it is 
the large pest, humans, that causes the greatest levels 
of concern. She notes the ordinary people are by and 
large, friendly and helpful. It is the bandits, police, 
_ military, industrialists and rioting natives that disrupt 
her life. 

As I started to read this book I became very dis- 
concerted. In the past few years I have looked for 
amphibians and reptiles in tropical jungles on many 
_ occasions. My success rate is very low. The author 
travels to the same locations in South America and 
immediately starts finding many individuals of multi- 
ple species. My estimation of my frog finding prowess 
dropped page by page. By the time the author reaches 
the 1990s, however, her success rate has plummeted. 
By 1996 she says “we hug and screech with excite- 
ment” on finding one frog. I realize that my lack of 
success is due less to my incompetence than to the 


Conversations with an Eagle 


By Brenda Cox. 2002. Greystone Books, Douglas & McIntyre 
Publishing Group, 2323 Quebec Street, Vancouver, Brit- 
ish Columbia VST 487. 261 pages, paperback. 


This is a story of dedication to an eagle rather than 
conversations with one. Brenda Cox was a volunteer 
worker at a raptor rehabilitation centre south of Van- 
couver (Orphaned Wild Life — OWL), and became 
enthralled with raptors of all kinds. Her special inter- 
est was a Bald Eagle, which was three months old 
when it arrived at the Centre, and her book describes 
the trials and tribulations of her efforts to train Ichabod 
so that she could be used in the OWL education pro- 
gramme. 

In the course of the story there are passages de- 
scribing the methods used to rehabilitate owls, hawks 
and eagles. When the eagle was still young its behav- 
iour was typically fearful, but the bird became more 
and more aggressive towards Cox, such that she had 
to stand against a pillar armed with a household mop 
to protect herself if she was attacked. In spite of the 
danger, she persisted in training the bird to come to 
her gloved arm, and eventually it obeyed three com- 
mands: Up, Wait and Off. But it was always uncer- 
tain whether the bird would cooperate or not on any 
given day. She enlisted the help of expert falconers in 
the Vancouver area. They were doubtful whether a 


Book REVIEWS 


rapid loss of frogs in my lifetime. This is even more 
depressing. When I go to Monteverde next spring I 
have little hope of seeing the fabulous Golden Frog. 
On 7 April 1987 this author saw “over one hundred 
dazzling bright golden toads“ at one small pool at 
Monteverde. 

The author also deserves another accolade for, while 
she 1s pursuing her scientific research, she is also be- 
ing a parent to two children. I was delighted with the 
way she involved them in her work and gave them 
opportunities to broaden their knowledge. This may 
have made her life harder and certainly caused her 
concerns, but I am sure these children will benefit. 

So if you want to find out about the life of a field 
researcher read this entertaining book. Follow the trials 
of first reaching the field locations using often unreli- 
able local transport. Join in meals that are good, bad, 
or bizarre. Meet local people, many of whom care 
greatly about their environment. Find out how a sci- 
entist collects data and uses it to create a new under- 
standing of our planet. 


Roy JOHN 


2193 Emard Crescent, Ottawa, Ontario, K1J 6K5 Canada 


Bald Eagle could be reliably trained, though they 
helped and advised her. Bald Eagles have seldom been 
trained successfully, while Golden Eagles have been 
used by falconers in many parts of the world. Cox 
was able to train the eagle such that she could take 
her from her cage to a perch in a field, using jesses 
and a stout leash attached to her waist, but there were 
increasing dangerous attacks which inflicted talon 
wounds on Cox’s head, arms and feet. 

After several years, the OWL management decided 
that they would never be able to use the bird in their 
programme and asked Cox to remove it from the facil- 
ity. In its new quarters, shortly afterwards the bird 
developed a lung disease and was euthanized. Cox’s 
dedication to her volunteer job is remarkable since, in 
the course of the seven years, she did not have steady 
employment or reliable housing or transportation. 
Since Ichabod’s death, she has now found a steady 
career: as a conductor with British Columbia Rail, 
and watches the wild raptors along the railway lines. 
Her story would appeal to someone interested in 
raptor rehabilitation methods and the lore of falconry. 


JANE ATKINSON 


255 Malcolm Circle, Dorval, Quebec H9S 1T6 Canada 


284 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


North American Owls, Biology and Natural History, Second Edition 


Paul A. Johnsgard. 2002. Smithsonian Institution Press, Wash- 
ington. 298 pages, 42 colour plates, 69 multi-figured black- 
and-white maps and sketches, 10 tables, 2 graphs. $46.25 
Canadian, hardcover. US$27.95. 


Although Dr. Johnsgard’s long history of produc- 
ing books about birds is well known, his entry to the 
“owl-lover’s camp” is more recent. The first edition of 
this book came out in 1988 and immediately became 
the standard reference book for owls of the United 
States and Canada. For the second edition, 14 years 
later, Johnsgard has added twelve species of Mexican 
owls and nearly doubled the number of references 
(although, contrary to convention, many of the recent 
references are not mentioned by name in the text!). 
Extremely few changes have been made to the excel- 
lent introductory chapters, but this leaves them less up- 
to-date than the individual species accounts. Often he 
cites the authors of the Birds of North America account, 
rather than the original observers who merit the credit; 
an example is Gottfred and Gottfred’s new informa- 
tion concerning courtship and copulation of the Great 
Horned Owl, published in Blue Jay in 1996. 

The strengths of the first edition have been main- 
tained: the introductory chapters and species accounts 
are neatly arranged; maps of each species’ range are 
detailed; Johnsgard’s pen-and-ink sketches of aspects 
of behaviour are delightful; the colour plates and pho- 
tographs are superb. Johnsgard writes well, but in a 
few occasions new material has been interjected clum- 
sily, marring the previous smooth flow of information 
in the first edition. 


Some compromises were necessary. To make room 
for the twelve Mexican species, detailed descriptions 
of plumage have been omitted in this second edition. 
This does not excuse three sloppy errors. Johnsgard 
tells of the Pygmy Owl range extending altitudinally 
to 37 000 meters in Mexico, whereas the quoted source 
gave a credible 3700 m. A paper by Jack Holt on Great 
Horned Owls in the Cincinnati region is credited to 
Denver Holt. Thirdly, when Johnsgard withdrew his 
overdrawn sketch of the “false eye-spots” on the back 
of the Northern Pygmy Owl’s head from the first edi- 
tion and substituted a more realistic sketch of the Fer- 
ruginous Pygmy Owl in the second edition, he failed 
to change the figure numbers in the text to comply 
with this change. 

These minor caveats aside, this masterful second 
edition will be welcomed by owl enthusiasts around 
the world, and should be purchased even by those own- 
ing the first edition. It is written more for the scientist 
than the amateur, but, by identifying deficiencies in 
our current knowledge, this book should offer possi- 
bilities for future study by graduate students and others 
who love our night-time friends. 

Johnsgard’s exquisite sketches, 10 coloured paint- 
ings of owls by Louis Agassiz Fuertes, and 31 fine col- 
our photographs, including nine of Mexican owls, make 
this an unusually attractive book. Buy it! 


MARTEN J. STOFFEL 


Box 183, RR#4, Saskatoon, Saskatchewan S7K 3J7 Canada 


The Mountain White-Crowned Sparrow: Migration and Reproduction at High Altitude 


By Martin L. Morton. 2002. Studies in Avian Biology 
Number 24. Cooper Ornithological Society, Camarillo, 
California. 236 pages, 8 colour plates, pencil illustrations 
heading each chapter. U.S.$27. 


The northern limit of the distribution of the Moun- 
tain White-Crowned Sparrow (Zonotrichia leucophrys 
oriantha) extends slightly into southern British Colum- 
bia, Alberta and Saskatchewan. Its main breeding range 
is in mountainous regions of the western U.S. Here 
in Waterton Lakes National Park we’re at the contact 
zone between oriantha and Z. |. gambelii. 

Morton’s 25-year study of oriantha in the Tioga 
Pass area of California’s Sierra Nevada ranks with P. 
J. Greenwood’s lengthy study of the Great Tit (Parus 
major) in Europe and G. E. Woofenden’s work with 
the Florida Scrub Jay (Aphelocoma coerulescens) in 
showing the value of continuous long-term field inves- 
tigations in advancing a broad spectrum of ideas and 
hypotheses in avian biology. Morton’s focus is differ- 
ent, however, in emphasizing physiology more than 
behaviour or ecology. In 1968, Morton recognized 


that there were significant gaps in knowledge of mi- 
gratory passerines on their summering grounds, espe- 
cially in mountains where large variations in environ- 
mental conditions occur. 

As my interests lie more with natural history, life 
history data (age at maturity, number, size and sex ratio 
of offspring, dispersal and survival rates) and ecolog- 
ical factors than with physiology, I found Morton’s 
first three chapters on Migration Arrival, Social System 
and Behavior, and Demography of most interest (al- 
though a bit disappointing at only one-quarter of the 
book’s pages). The dynamics of arrival of oriantha 
varied greatly from year-to-year depending upon the 
amount of remaining snowpack and the frequency of 
spring storms, as well as by age and sex. By trapping 
and colour-marking individuals, the researchers found 
that older males (age 2+ years) generally arrive ear- 
liest, followed by one-year-old males, older females 
and one-year-old females. Throughout the monograph, 
Morton does an excellent job of analysing observed 
data in terms of costs and benefits, and ecological fact- 


2004 


ors, in this case, suggesting that older, experienced 
birds knew the migration route and recognized the 
_ breeding area once they reached it, even it if was snow- 


covered. 

Morton’s efforts over many reproductive seasons 
made it possible to measure mate fidelity, age of mates, 
frequency of polygamous pairings, agressive behav- 
iours, and the functions of vocalizations, in addition 
to the usual study of territory establishment, pairing, 
and between-year breeding dispersal. 

Although I expected the chapters on Gonadal Con- 
dition, and Body Size and Body Condition, to be of 
less personal interest, Morton’s explanations of the 
connections between physiology and behaviour, and 


| description of the role of environmental cues in annual 
_ cycles made these sections much more interesting than 


anticipated. Environmental factors are either ultimate 
(e.g., availability of an adequate food supply, predation 
pressure, weather patterns) or proximate (e.g., photo- 


| period, ambient temperature) in their effects on the 
timing of reproduction. 


Nest history (chapters 7, 8, 9 and 10 on Nests and 
Eggs, Nestlings and Fledglings, Nest Failure and 


Book REVIEWS 


Reproductive Success, respectively) provided reliable 
information on physiological and behavioural respons- 
es of breeding birds to environmental variation, often 
to the level of individuals because of Morton’s use of 
marked birds. And the researchers determined that 
the snowpack, because of its effects on nesting sched- 
ules and nest locations, was a stronger environmental 
factor on reproductive success than sub-freezing tem- 
peratures or summer storms. Oriantha exhibited plas- 
ticity in responding to snow conditions by abandon- 
ing ground-nesting and building their nests in elevated 
sites when there was more snow, as opposed to Her- 
mit Thrushes (Catharus guttatus) or Dark-eyed Juncos 
(Junco hyemalis), which nested on the ground no 
matter what the environmental conditions. 

Morton’s study shows that challenges posed by en- 
vironmental variation often can be met with existing 
behavioural and physiological responses; adaptation 
occurs through flexibility rather than through acqui- 
sition of new abilities or mechanisms. 


CynpI M. SMITH 
Box 5, Waterton Park, Alberta TOK 2MO Canada 


Seabird Bycatch: Trends, Roadblocks, and Solutions 


Edited by Edward F. Melvin and Julia K. Parrish. 2001. Uni- 
versity of Alaska Sea Grant AK-SG-01-01, Fairbanks, 
Alaska. viii + 206 pages. U.S.$20.00. 


The incidental catch of various water birds and other 
aquatic organisms (e.g., turtles, sea otters, porpoises, 
seals) in fishing nets has long been of concern to 


naturalists for conservation reasons and to the fishing 


industry for economic reasons. However, until recent- 
ly this issue has been on the “back burner” compared 
with more pressing and widespread environmental 
problems. The advent of intensified fishing efforts, 
huge factory ships, and collapsing fish stocks have 
elevated this bycatch into both a major international 
environmental issue and a serious economic problem. 
This book, essentially the proceedings of a sympo- 
sium, presents a recent update on the extent of the 
problem in marine waters, world-wide, evaluations 
of several proposed solutions as they apply to specific 
fisheries and problems that have yet to be overcome. 
The editors begin the book with a short preface that 
outlines the history and extent of the problem, espe- 
cially since it was identified as a major conservation 
issue in the early 1970s. The first full chapter is a syn- 
thesis, also by the editors, of a symposium organized 
by the Pacific Seabird Group in Blaine, Washington, 
in 1999. The synthesis summarizes the complexity of 
the problem, “roadblocks” in the way of solving it and 
guidelines to possible solutions. The remainder of the 
book consists of nine peer-reviewed, scientific papers 
(eight of which were presented at the symposium) 
and seven abstracts. Two of the papers were reprinted 
from recent (1999 and 2000) scientific journals. One 


paper, by John Cooper, John Croxall and Kim Rivera 
outlines the complexities of international efforts to re- 
duce bycatch through research, local and international 
regulations and cooperation among scientists, the fish- 
ing industry and conservation groups. An afterword by 
Craig Harrison graphically illustrates the complexity 
of the problem by outlining the political and practical 
difficulties of coordinating efforts to reduce bycatch 
in the sockeye salmon fishery in the “shared waters” 
of British Columbia and Washington, where two fed- 
eral, one provincial and one state government share 
jurisdiction with “21 Native American tribes” [a total 
that applies only to Washington and omits the first 
nations of British Columbia]. 

The rest of the papers and abstracts document the 
extent of the bycatch in specific fisheries, results of 
various experiments to reduce bycatch, and attempts 
to measure the impact of specific fisheries on specific 
populations of birds. As some seabirds spend non- 
breeding times thousands of kilometers from their 
breeding sites, measuring the effects of a specific fish- 
ery on a particular breeding population is very diffi- 
cult, especially as many seabirds are long-lived and 
don’t breed every year. Moreover, fishing in a given 
area may affect some species much more profoundly 
than their close relatives (for example, effects of by- 
catch on Black-footed Albatross populations are much 
more significant than on Laysan Albatrosses) and may 
affect different age groups differently (for example, 
juvenile Black-footed Albatrosses are caught at a high- 
er rate than adults). Research presented in this vol- 
ume from the Atlantic, Pacific and Southern (Antarc- 


286 


tic) oceans shows that modifications to fishing gear 
(weights, design, visual and acoustic alerts), timing 
of fishing (daily and seasonally) and other techniques 
can reduce bycatch substantially without substantially 
reducing the take of target species, but that these mod- 
ifications have different effects on the rate of bycatch 
of different species and sometimes of a given species 
in different areas, different times of the day or different 
seasons. Banding continues to be helpful in sorting 
out the origins of birds caught in specific fisheries, 
while higher tech tracking devices and research on 
genetic markers are also starting to improve our knowl- 
edge of the complexities of movements by these highly 
mobile species. Albatrosses receive the greatest amount 
of attention, but data on various other “tubenoses,” 
alcids, larids and some marine mammals are also pre- 
sented. 

Canadian content figures prominently in this book. 
A 1972 paper by Canadian C. Eric Tull and co-authors 
on the magnitude of Thick-billed Murre mortality in 
Greenland fisheries is credited (page v) in drawing 
attention to the significance of fisheries in seabird 
population declines, and international legal efforts to 
reduce bycatch in longlines stem from a resolution at 
a 1996 meeting in Montreal of the World Conserva- 
tion Congress of the World Conservation Union (page 
11). Data from British Columbia are included in a 
genetics (DNA) study of the breeding locations of 
Common Murres entangled in fishing nets in Wash- 
ington’s San Juan Islands and Puget Sound (pages 
125-127) and in an abstract (pages 191-192) on sea- 
bird avoidance experiments in northern Pacific waters. 
Other papers and abstracts on observations and re- 
search in Alaskan and Washington waters involve 
populations of birds that spend portions of their lives 
in British Columbian waters. Research from Atlantic 
coast provinces and British Columbia are cited fre- 


The Life of Mammals 


By David Attenborough. 2002. Princeton University Press, 
New Jersey. 320 pages, U.S.$29.95. 


As the latest edition in the BBC “Life” series com- 
bining television and print media, the book on mam- 
mals is nicely illustrated with a good selection of 
colour photographs that closely follows the engaging 
text by David Attenborough as he presents interesting 
stories on these fascinating animals. The first chapter, 
“A Winning Design’, starts off with the ability of 
mammals to adapt to different environments on earth, 
including the harsh arctic conditions where lemmings 
live year round. After describing some basic charac- 
ters of mammals, such as hair and the production of 
milk, there is a general introduction to the origin and 
evolution of this group of warm-blooded organisms. 
The chapter finishes off with two early mammalian 
radiations that cover the egg-laying monotremes and 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


quently. 

This book provides a useful compilation of recent 
developments on a complex issue, with plenty of in- 
formation on legal (national and international)/poli- 
tical aspects, recent research on seabird movements, 
recent research on genetics of seabird populations, and 
practical tests of various techniques under a variety of 
conditions in a variety of waters. The literature cited 
sections provide numerous additional references for 
those requiring further information. The book should 
be on the library shelves of any researchers, govern- 
ment agencies and non-government conservation and 
scientific organizations studying seabird biology and/ 
or the conservation of seabirds. Errors appear to be 
few — the reference to Wilson et al. (1985) cited on 
page 116 is not listed in that chapter’s literature cited 
section, which includes two references that don’t appear 
to have been cited. However, since most of tne chapters 
are written as scientific papers, some may be too tech- 
nical for some readers. The review by Cooper et al. 
of legal and quasilegal aspects of the issue provides a 
valuable reference compendium, but is so riddled with 
acronyms that I found myself wishing for a glossary. 
I also hope that the seven abstracts at the end of the 
book are expanded into full papers somewhere. As 
valuable a contribution as the book is in itself, this 
volume will undoubtedly also stimulate more research 
that will require another update before long. Such an 
update could usefully also include papers or chapters 
on aquatic bird bycatch of fisheries on inland waters, 
such as those of the Great Lakes, several large prairie 
province lakes and similar lakes on other continents. 


MARTIN K. MCNICHOLL 


4735 Canada Way, Burnaby, British Columbia V5G 1L3 
Canada 


the marsupials, which give birth to under-developed 
young. 

The remaining nine chapters deal with the placen- 
tal eutherian mammals but instead of continuing to 
describe them by scientific groups the format switches 
to artificial categories such as diet and habitat. This 
unnatural classification seems awkward at times with 
bats, which people can readily identify with, split into 
two separate chapters, “Insect Hunters” (curiously 
including vampire bats) and “Life in the Trees” (al- 
though some insect-eating bats also live in trees). There 
was a missed opportunity to educate readers in scien- 
tific classification and evolution while still entertaining 
them with a plethora of amazing natural history stories. 

Primates get star billing in the book with the last two 
and a half chapters devoted to this charismatic order 
of mammals. Although the higher-level taxonomy (or 
common names employed) is not current, it begins 


2004 


_ with prosimians as an early branch of the primates. 
The latest view is that this is not a natural group be- 
cause, for example, tarsiers are more closely related 
_to monkeys and apes, as alluded to but nonetheless 
still included in the chapter with lemurs. The other 
primates are grouped to cover the new and old world 
_ monkeys, and ending with the gibbons and great apes, 
including a branch for humans. The last half chapter 
concentrates on both physical and social anthropology 
from the first evidence of bipedal locomotion to cul- 
tivation and civilization. 
The book is definitely aimed at a general but knowl- 
_ edgeable audience with an interest in nature and mam- 
_mals. I am sure, however, that practising biologists 
will still find a few facts new to them in their non- 
specialist group because the background research is 
_ relatively good. It was nice to see some recent scien- 
tific hypotheses on mammalian evolution making it 
into the book such as the close relationship between 


BOOK REVIEWS 


whales and hippopotamuses. But some other emerging 
ideas based primarily on molecular data did not, in- 
cluding the association of bats with carnivores, ungu- 
lates and whales, as opposed to insectivores, or tree 
shrews, flying lemurs and primates. 

My criticisms are mostly biologically oriented be- 
cause the book is attractively presented with most 
photographs of good quality and information well writ- 
ten. But there should be more books that combine 
current scientific research with an explanation of the 
deeper implications or processes involved for wider 
distribution to the general public looking for mean- 
ingful substance beyond the usual cursory facts. 


BurRTON K. Lim 
Centre for Biodiversity and Conservation Biology, Royal 


Ontario Museum, 100 Queen’s Park, Toronto, Ontario M5S 
2C6 Canada 


_ Geographic Variation in Size and Shape of Savannah Sparrows (Passerculus sandwichensis) 


_ By James D. Rising. 2001. Studies in Avian Biology Number 
23. Cooper Ornithological Society, Camarillo, California. 
65 pages. U.S.$7. 


The newer field guides, such as The Sibley Guide to 
Birds, occasionally describe and illustrate subspecies 
of birds, usually based on differences in plumage or 
_ soft parts. The “bander’s bible”, Peter Pyle’s Identi- 
_ fication Guide to North American Birds, includes 
wing and tail measurements to help banders identify 
birds to the subspecies level — for example, he dis- 
cusses 14 subspecies of the Savannah Sparrow. But 
have you ever wondered why these differences exist 
in the natural world? 

Rising does and asks two questions in this regard: 
Why do features such as body size, wing length, or bill 
size and shape differ across a species’ range? and, if 
these differences reflect adaptations to the different 
environments to which the species is exposed, what 
are the selective factors that have caused them? 

One of the classic explanations for geographic vari- 
ation in size is Bergmann’s Rule, which holds that 
individuals of a species (vertebrates only) from colder 
areas are generally larger-bodied than individuals from 
warmer areas. Allen’s Rule takes this one step fur- 
ther, stating that within such species, individuals from 
colder areas will have smaller appendages relative to 
their body size than individuals from warmer areas. 

Rising asked his questions of the Savannah Spar- 
row, which is one of the most wide-spread songbirds 
in North America. He describes and quantifies geogra- 
phic variation in the species throughout its breeding 
range, from Alaska to the Maritimes to central Mexico, 
and relates trends in phenotypic variation to environ- 
mental variation. 

He found some clinal variation in size of Savannah 
Sparrows, with birds from the northeast being slightly 
larger than those from the west, and birds in cool, 


moist areas were larger than those where it is hot and 
dry. But the species overall did not seem to follow 
Bergmann’s Rule; rather, measures of summer tem- 
perature and precipitation explained well the patterns 
of size variation. His more significant find, though, 
was that birds were larger on islands than on main- 
land sites, whether in the Aleutian Islands, Alaska, or 
on Sable Island, Nova Scotia. He speculates that the 
long, cool, moist summers on these islands results in 
a predictable and fairly rich food supply. This, com- 
bined with the rather long breeding season, allows 
multiple broods and perhaps enhanced competition 
for high quality territories. This competition for either 
food or territories might select for larger body size. 

Rising ends with some taxonomic comments, com- 
ing out on the side of the “lumpers’’. He sees no virtue 
in naming subspecies where the only way they can be 
reliably separated is by locality. He suggests recog- 
nizing only two subspecies of non-saltmarsh Savannah 
Sparrows, P. s. sandwichensis (large size) and P. s. 
princeps (large and pallid), whereas the nine saltmarsh 
subspecies seem to be clearly separable by morpho- 
logical characters. This is in contrast to the 17 sub- 
species currently recognized by the American Orni- 
thologists’ Union. 

Rising’s work relies on Principal Components and 
Discriminant Functions analyses, which, although I 
found it a little heavy going, was well presented sequen- 
tially and a good example of the use of these techniques 
for morphometric comparisons. 

It will be interesting to watch in the coming years 
how Rising’s thorough morphological studies interact 
with genetic analyses in assessing variation and the sub- 
species of Savannah Sparrows. 


CyNnbDI M. SMITH 


Box 5, Waterton Park, Alberta TOK 2MO Canada 


288 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Warblers of the Great Lakes and Eastern North America 


By Chris Early. 2003. Firefly Books Ltd., 3680 Victoria 
Park Avenue, Toronto, Ontario, M2H 3K1. 131 + pages. 
Cloth $24.95: paper $16.95 


Sparrows and Finches of the Great Lakes and Eastern North America 


By Chris Early. 2003. Firefly Books Ltd., 3680 Victoria 
Park Avenue, Toronto, Ontario, M2H 3K1. 128 + pages. 
Cloth $24.95: paper $.16.95 


These books are an expansion and revision of two 
earlier books by Chris Early (Warblers of Ontario and 
Sparrows and Finches of Ontario). They both follow 
the successful format of the earlier books. Each spe- 
cies is shown in two to five photographs (Warblers) 
or two to seven photographs (Sparrows). The actual 
number of photographs for each species depends on 
how variable the bird’s plumage can be. A short intro- 
ductory note is followed by descriptions of the key 
plumage characteristics and other relevant details. A 
5 x 3 cm map of America shows the summer and 
winter ranges. 

“Warblers” covers 37 species plus one hybrid in 
some detail. An additional seven vagrant species and 
one race (the Yellow-rumped “Audubon’s” Warbler) 
are covered by brief comments and a single accom- 
panying photograph. 

“Sparrows” includes 25 species covered in full and 
20 vagrant species. In addition to the North American 
sparrows (buntings) this volume includes longspurs, 
finches, grosbeaks, crossbills, snow bunting and house 
sparrow. 

There are two features used by this author that will 
make this book particularly valuable to beginners. 
The first is called a cheat sheet and lists all the birds 
in groups according to a key characteristic — such as 
an unstreaked breast. The birds are further grouped 
by a second characteristic such as a breast spot. The 
second, and even better innovation, is a series of com- 


Birds of the Yukon Territory 


Edited by P. H. Sinclair, W. A. Nixon, C. D. Eckert and N. 
L. Hughes. University of British Columbia Press, Van- 
couver, British Columbia. 


My appetite for a long-overdue return trip to the 
Yukon has been incredibly whetted! This book is not 
just an atlas, but also a coffee table book, a very use- 
ful bird reference, and a source necessary for a 
planning a trip to the Yukon. This book has stunning 
photographs throughout, and not just of birds, but of 
dramatic scenery (i.e., habitat shots) that will make 
birders and non-birders alike think, “I must go there.” 

Prior to the species accounts, there are chapters 
covering the environment, aboriginal use of birds, 
conservation, history and a month-by-month overview 
of birdlife. There is also a short chapter outlining the 


parison tables. These consist of a set of photographs 
with birds in similar poses. The page is arranged to 
show the most similar birds close together. For example, 
there are three Common Redpoll photographs imme- 
diately above three equivalent Hoary Redpoll photo- 
graphs. Similarly, all the rufous-capped sparrows are 
on one page. I think the arrangement of the warbler 
comparison table is very clever. Here the Mourning 
Warbler photograph is adjacent to a Connecticut War- 
bler photograph. In turn, the Connecticut is above a 
photograph of a Nashville Warbler, which in turn is 
above a Northern Parula. All four birds share similar 
combinations of colour and cause confusion to the 
novice. This arrangement allows for easy compari- 
son, either up and down or across the rows. For the 
warblers there are tables showing the spring and fall 
plumages. These features are on fold-out sheets at the 
end of the book, making them quickly available in the 
field. At the front of the book is a seasonal distribution 
list for Point Pelee National Park. Using line thickness 
the author also indicates the abundance of each species. 
I find such distribution lists to be extremely useful. 
These then are two small, portable books that deal 
with birds that many find difficult to identify. The text 
is well written. The photographs are both beautiful and 
illustrative. Any one who needs help with the species 
covered will find these books a tremendous help. Per- 
haps we can encourage the author to write a third 


book on shorebirds. 
Roy JOHN 


2193 Emard Crescent, Ontario, K1J 6K5 Canada 


data management for the more than 160 000 records 
that were used. 

Each species account spans one to three pages. All 
species are illustrated by a line drawing and at least 
one, sometimes several, colour photographs; frequent- 
ly, there is a very good habitat shot as well. Each 
account also has a distribution map, and a histogram 
outlining the number of database records for each 
week of the year. The base layers of the maps show 
major roads and rivers as well as ecozones; I would 
have liked to see the dominant population centers 
marked as well though. 

The text is thorough and reads well. All the expect- 
ed sections are there (distribution, nesting, habitat...), 
and, where appropriate, a section on aboriginal use of 


2004 


the bird is included. Several appendices and a 600+ 
reference list end the book. 

The items that bothered me were few, and mostly 
minor. Noteworthy was the absence of birding essays, 
a feature I first saw in Birds of Delaware, and thought 
added spark to the thick reference book. Page numbers 
were located in the most irritating choice possible, 
along the inner margin of each page, thus not permit- 
ting a rapid flip-through while searching through this 
600 page book. The final, and dare I say incompre- 
hensible feature of the book was the uninteresting and 
out of focus (soft focus?) cover shot of the Territorial 
bird, the Common Raven. People do judge a book by 


BOTANY 


BooK REVIEWS 


289 


its cover, rightly or wrongly, and in a book with hun- 
dreds of stellar shots, I remain baffled as to why that 
particular shot was chosen to be most people’s first 
impression of the book. 

Overall though, this is a very, very good book, and 
a must-have for people interested in the birdlife of 


the North. 
RANDY LAUFF 


Department of Biology, St. Francis Xavier University, Anti- 
gonish, Nova Scotia B2G 2W5 
Literature Cited 


Hess, G. K., R. L. West, M. V. Barnhill I, and L. M. Fleming, 1999, 
Birds of Delaware. University of Pittsburgh Press. 635 pages. 


Wild Flowers of the Yukon, Alaska & Northwestern Canada, 2™ Edition 


By John G. Trelawny. 2003. Harbour Publishing Co. Ltd., 
P.O. Box 219, Madeira Park, British Columbia VON 2HO 
Canada. 224 pages. $24.95. 


The first edition of this wild flower book was pub- 
lished in 1983 and I reviewed it in 1984. The layout 
in this new volume is much better. The type is larger 
and more spaced on a purely white paper and is 
wrapped around the absolutely beautiful colour pho- 
tographs. 

Many of the pictures of the 332 species treated are 
the same as those in the first edition but the more 
modern reproductions and paper have made them 
shine. The descriptive text for each species has been 


only slightly revised. The surrounding text includes 
Acknowledgments, an easy-to-read map, an Introduc- 
tion, an Illustrated Glossary, a Key to Species using 
Flower Colour and Shape, Photo Credits, Bibliogra- 
phy, an Index, and a list of Additional Field Guides 
from Harbour Publishing. This is a most interesting 
and delightful book for anyone living in, visiting or 
planning to visit this wonderful area. 


WILLIAM J. Coby 


Biodiversity, Program on Environmental Health, Agriculture 
and Agri-Food Canada, Wm. Saunders Building, Central 
Experimental Farm, Ottawa, Ontario K1A 0C6 Canada. 


Wild Flowers of Field & Slope in the Pacific Northwest 


By Lewis J. Clark. 2002. Harbour Publishing, P.O. Box 
219, Madeira Park, British Columbia VON 2HO. $9.95. 


Wild Flowers of Forest & Woodland in the Pacific Northwest 


By Lewis J. Clark. 2003. Harbour Publishing, P.O. Box 


219, Madeira Park, British Columbia VON 2H0. $12.95 


Wild Flowers of the Mountains in the Pacific Northwest 


By Lewis J. Clark. 2003. Harbour Publishing, P.O. Box 
219, Madeira Park, British Columbia VON 2HO0. $12.95. 


These three books contain absolutely beautiful col- 
our photographs. The photographs are numbered 
sequentially and each one has a marker [x0.5] to in- 
dicate its size. Each is accompanied by a printed para- 
graph with the common and scientific names, a de- 
tailed description, together with the habitat and range, 
and a number to indicate its sequence in the book. 
There are 108 in the first book, 100 in the second and 
106 in the third. 

There is a four- or five-page interesting introduction 
at the front of each book and an index, glossary and a 


list of Additional Field Guides available from Har- 
bour Publishing at the end. There are most interest- 
ing pictures on the front covers of each book. On the 
back covers there is a note about the author, Dr. Lewis 
J. Clark, together with a map of the Pacific Northwest 
on which there are shaded areas depicting where the 
wild flowers can be found. All are elegant. 


WILLIAM J. Copy 
Biodiversity, Program on Environmental Health, Agriculture 


and Agri-Food Canada, Wm. Saunders Building, Central 
Experimental Farm, Ottawa, Ontario K1A 0C6 Canada 


290 


MISCELLANEOUS 


The Canoe: A Living Tradition 
By John Jennings. Firefly Books, Toronto. 272 pages. $59.95. 


There could hardly be a more poetic way to ex- 
plore the natural world than by canoe. While not all 
observers of natural history are canoeists, most canoe- 
ists have respect for and curiosity about nature. In- 
deed, a loon’s call or the slap of a beaver’s tail, accom- 
panied by the quiet burbling of a canoe’s wake, has 
served as an introduction to the contemplation of 
nature for countless North Americans. 

The Canoe: A Living Tradition is, in a sense, a nat- 
ural history of traditional North American watercraft. 
Included in this beautifully illustrated, coffee table- 
sized book are chapters dedicated to the history and 
construction of birch bark and dugout canoes, as well 
as skin kayaks and umiaks (large, open vessels from 
the Canadian Arctic). This book also discusses the 
influence of canoes on the fur trade and the settlement 
of northern North America, as well as the develop- 
ment of recreational canoes and the preservation of 
canoe history in more recent times. A respected expert 
writes each chapter on the type of craft or historical 
context being discussed. This makes for inconsistent 
writing at times, but also gives the book an authority 
that would not have been achieved by a single author. 
I found that the technical details about the dimen- 
sions (i.e., width and length, etc.) of canoes a bit repet- 
itive. On the other hand, the documentation of these 
measurements may be valuable in the future. As John 
Jennings notes in the first chapter, the original canoe 
designs of several First Nations in North America have 
been erased without a trace, a poignant reminder of 
the terrible loss of traditional knowledge that contin- 
ues to this day. 

While the book brilliantly covers the historical 
context of canoeing and canoe building (over the 
span of 230 pages), it pays scant attention to modern 
uses of the canoe (2 pages). After all, as the title sug- 
gests, canoeing and canoe building is a “living tradi- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


tion’. Canoeing is still one of the most elegant ways to 
explore nature and there are many fine canoe builders 
in North America currently manufacturing modern 
and traditional canoes from a wide range of natural 
and synthetic materials. It seems as though a chapter 
could have been devoted to recent developments in 
the design and construction of modern canoes. 

The book gives a good overview of how birch bark 
and dugout canoes are constructed, including useful 
photographic illustrations; however, before you run 
out and start assembling materials, take heed: the 
information provided is insufficient for the “do it your- 
self’ canoe builder. A noticeable shortcoming is the 
omission of information about the many resources 
available to those interested in building traditional 
birch bark and dugout canoes, as well as skin kayaks. 
For instance, while the book contains a photograph 
of a canoe built by César Nawashish, it fails to refer- 
ence the excellent 1971 National Film Board film 
“César’s Bark Canoe”, which documents Nawashish’s 
completion of a bark canoe with exquisite detail. 
Other contemporary builders of traditional craft have 
also produced thorough “how to” publications on 
building birch bark canoes and skin kayaks including 
David Gidmark, Robert Morris and Wolfgang Brink, 
which have not been referenced in this book. 

While Adney and Chapelle’s Bark Canoes and 
Skin Boats of North America will remain the bible of 
traditional canoe and kayak design, The Canoe: A 
Living Tradition is broader (literally and figuratively), 
more colourful and more accessible. It is simply irre- 
sistible for the canoe enthusiast. Its shortcomings are 
few and its design and production qualities are excel- 
lent. In summary, it is a valuable addition to the annals 
of canoe culture. Congratulations to John Jennings 
and the Canadian Canoe Museum for their fine work. 


PATRICK WILLISTON 
41 Nielson Road, Smithers, British Columbia VOJ 2N2 Canada 


A Passion for Wildlife: The History of the Canadian Wildlife Service 


By J. A. Burnett. UBC Press, Vancouver, British Columbia. 
Hardcover. 331 pages, $27.95 paper, $85.00 cloth. 


Anyone who is prejudiced against “civil servants” 
should read this book. The Canadian Wildlife Service, 
from its inception, has contained a cadre of dedicated 
scientists who work long hours, share their knowledge 
widely, and on occasion risk their lives for the sake 
of the environment. Burnett, an insider, tells the story 
of this valuable and generally cost-effective organiza- 
tion, beginning in 1916 with the five men who were 
involved in drafting and implementing, from the 
Canadian side, the Migratory Birds Convention Act. 
In 1918, Hoyes Lloyd was appointed as Supervisor 


of Wild Life Protection in Canada. Lloyd, in turn, 
appointed three Federal Migratory Bird Officers. The 
Dominion Wildlife Service (now the Canadian Wild- 
life Service or CWS) was founded on | November 
1947. 

Burnett follows a roughly chronological sequence, 
with 67 photographs. His writing skills introduce us 
to many interesting scientists and their achievements, 
without the inevitable plethora of names becoming 
objectionable. The footnotes list the main publica- 
tions that resulted from extensive research. He covers 
objectively the lowest point in CWS history, the short 
but destructive reign of Suzanne Blais-Grenier as 
federal Minister of the Environment. 


2004 


This excellent history will be of interest to most 
ornithologists, mammalogists and environmentalists, 
and will be a useful reference in every major library. 

However, I must admit to being flabbergasted by 
the publication of this book by a reputable university 
press, without any mention, let alone acknowledge- 
ment, of the previous publication, with the identical 
title, and with 99 per cent of the text which filled one 
issue of The Canadian Field-Naturalist 113:1-214, 
January-March 1999. Readers of this journal do not 
need to buy the book; they already have a soft-cover 
version on their shelves. 

What does the new version offer as compared to 
the unmentioned and unacknowledged version? An 
appropriate foreword by Janet Foster and a few new 
photographs have been added, but most are unchanged 
except that their quality has been improved and some 
have increased in size. Additions have been made to 
the following sub-chapters: “Seabirds,” a paragraph 
about Tony Gaston’s work on the Ancient Murrelet; 
“Shorebirds,” two paragraphs on the Latin American 
cooperation in shorebird banding and mention of the 
Kees Vermeer Award now presented annually at Simon 
Fraser University; “Habitat Management,’ a paragraph 
on the purchase of Westham Island in the Fraser 
River delta; “Consolidation,” two new paragraphs on 
wildlife policy changes; “Peregrine Falcon,” Richard 
Fyfe’s investiture into the Order of Canada on 26 
April 2000; “Species Protection,’ a paragraph on the 
delays and rocky road of Bill C-5, the Canada Species 
at Risk Act, which finally passed on 11 June 2002; 
“A Work in Progress,” four sentences of reminiscences 


Birding on Borrowed Time 


By Phoebe Snetsinger. 2003. ABA Inc. Box 6599, Colorado 


Springs, Colorado. 307 pages. Paper US$19.19. 


I never met Phoebe Snetsinger. In fact I have only a 
vague recollection of hearing her name. I was sur- 
prised at her sudden prominence after she died. Now 
I have read her book and I understand. 

I expected someone who saw over 8000 species to 
be the typical “combat “ birder; a person who relishes 
getting the tick on her checklist above everything 
else. Phoebe Snetsinger was not a combat birder; she 
was a true birdwatcher. She insisted on really seeing 
and recording the birds she counted. She did not in- 
clude those flits that zip into view for an instant and 
are identified by the local guide. I have seen combat 
birders joyfully tick off such birds on their list and 
immediately move rapidly on in search of their next 
“lifer.” Indeed the author did not seem to be truly 
competitive until after her 7000" bird species and in 
line for the Guinness Book of Records. She ceased 
being competitive after achieving the world record at 
her 8000" species of bird. 

Her book is the account of her travels in accumu- 
lating this incredible total, and the joy she had in see- 


BOOK REVIEWS 


29] 


by Rob Butler. The title of Chapter 5 has been im- 
proved from “Policy Implementation” to “Emergence 
of Environment Canada.” A few small editorial im- 
provements are evident, such as a change from “at the 
same time that” to “even as.” Otherwise the two ver- 
sions are almost word for word and paragraph for para- 
graph in the text and word for word in the index. Sadly, 
an opportunity to index each annual wildlife conference 
was not seized. 

What has been deleted? A short biography of the 
author, Sandy Burnett; a short afterword by editor, 
Francis Cook, explaining the contributions to editing 
of the Canadian Field-Naturalist made by Harrison 
F. Lewis, C. H. D. Clarke, and A. J. Erskine; and a 
useful 31-page bibliography of major publications by 
CWS personnel, compiled by Erskine. 

The Canadian Field-Naturalist exists to record 
and spread information of value, and contains a clear 
statement that copyright for this particular paper re- 
sides with “Her Majesty the Queen in Right of Canada” 
as stated in 113(1). However, I remain surprised that 
the author and the publisher appear, consciously or 
unconsciously, to have chosen to withhold information 
about this material having previously appeared in the 
very scientific journal: in which this review is now 
appearing. 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N OJ8 
Canada 3 


Copies of CFN 113(1) are available for $10.00 from the 
Business Manager. 


ing all of these birds. She was a focused, energetic and 
hardworking individual who took her birdwatching 
extremely seriously. She studied her target species long 
before trying to see them. She kept copious notes on 
what she saw. I agree with this approach and try to 
use it myself. I do not understand why people pay so 
much money and effort to go to a prime nature locale, 
yet do not do their homework. 

Spurred into action by the threat of cancer and the 
potential to die young, Phoebe Snetsinger began a life 
of traveling for birds. In the process she made numer- 
ous friends in over 60 countries. I estimate she did ten 
to a dozen trips a year and spent about U.S.$50 000/ 
year in today’s currency. Overall, she must have made 
hundreds of trips. 

A neat bonus is the group of 29 black-and-white 
and 16 colour illustrations by Douglas Pratt. They por- 
tray the key milestone birds of Phoebe’s impressive 
odyssey. They are well done and make a nice coun- 
terpoint to the text. 

This is certainly a book for all birders. However, it 
is not simply a long list of birds seen, but a rampag- 
ing account of Phoebe’s obsession. This lady survived 


297 


gang rape, robbery, accidents, family crises, mechani- 
cal breakdowns, bad service and bad food against a 
background threat of cancer. Nothing seemed to slow 
her down. It is a tale of the fortitude of the human 
spirit. Indeed, I think a revision of this with the birding- 
related material severely edited (after all, which non- 
birder will realize the importance of a Red-shouldered 
Vanga — a small, House Sparrow-sized grey, black- 
and-white bird with bright chestnut shoulders — and 


New TITLES 


Zoology 


+The American Bison. By D. Lott. 2004. University of 
California Press. 229 pages, $40 U.S. Paper. 


+ Annotated Bibliography of the Quaternary Vertebrates 
of Northern North America. 2003. Edited by C. R. Harring- 
ton. University of Toronto Press. xxii plus 539 pages, not 
illustrated. $150 Cloth, $75. Paper. 


Annotated Checklist of the Birds of Chile. By Manuel 
Marin. Lynx Edicions, Barcelona, Spain. US$15.75. 


Behavior and Ecology of Pacific Salmon and Trout. By 
T. Quinn. UBC Press, 2029 West Mall, Vancouver, British 
Columbia V6T 1Z2 Canada. 400 pages, $85. 


Big Cat Diary: Leopard. By Jonathan Scott and Angela 
Scott. Harper Collins. 128 pages, $42. 


* The Bird Almanac. By D. Bird. 2004. Key Porter Books, 
70 The Esplanade, Toronto, Ontario, Canada. xvii + 460 
pages, not illustrated, $24.95. Paper. 


Birding in Venezuela. By Mary Lou Goodwin. Lynx Edi- 
cions, Barcelona, Spain. US$ 5.60. 


Birds of South Asia: The Ripley Guide. Volumes I and II. 
By Pamela C. Rasmussen and John C. Anderton. Lynx Edi- 
cions, Barcelona, Spain. US$95. 


Bowerbirds — Ptilonorhychidae. Edited by Clifford Frith. 
Oxford University Press, 70 Wynford Drive, Don Mills, 
Ontario M3C 1J9 Canada. 416 pages. $209 Cloth. 


+ British Columbia - A Natural History. By R. and S. 
Cannings. 2004. Greystone Books, Douglas and McIntyre, 
Vancouver, British Columbia, Canada. ix + 341 pages. 
$39.95 Paper. 


+ The Buffalo Wolf. 2003. By Lu Carbyn. Smithsonian 
books, Washington, D.C., USA. 248 pages. Paper. 


Curassows and Related Birds. By Jean Delacour and Dean 
Amadon. Update Chapter by Josep del Hoyo and Anna 
Motis. Illustrated by Albert Earl Gilbert. Lynx Edicions, 
Barcelona, Spain. US$75. 


7 Ecology and Conservation of Birds in the Salton Sink: 
an Endangered Ecosystem. By W. David Shuford and Kathy 
C. Molina. 2004. Studies in Avian Biology Number 27. 
Cooper Ornithological Society. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


its ilk?) would make it ideal for one of those inspira- 
tional stories we find in Reader’s Digest. Having final- 
ly seen a Red-shouldered Vanga, Phoebe was asleep in 
the tour bus when the bus crashed and she was killed. 
So she did not die of cancer, but binoculars in hand, 
on a birding trip. I never did meet this lady — pity! 


Roy JOHN 
2193 Emard Crescent, Ottawa, Ontario K1J 6K5 Canada 


* A Field Guide to the Bird’s Nests and Eggs of Alaska 
Coastal Tundra. By T. Bowman. 2004. Alaska Sea Grant 
College Program. 


+ A Guide to Hawk Watching in NA. By D. Heintzleman. 
2004. The Globe Pequot Press. 


+ Mammal Tracks and Signs. By Mark Elbroch. 2004. 
Stackpole Books. 


+ Key Marine Habitat Sites for Migratory Birds in Nuna- 
vut and the Northwest Territories. 2004. By M. Mallory 
and A. Fontaine. 2004. Canadian Wildlife Service, Ottawa, 
Ontario K1A OH3. Paper. 


7 Orca — Visions of the Killer Whale. By P. Knudtson. 
2004. Greystone Books, Vancouver, British Columbia. xvii 
+110 pages. $19.95. Paper. 


Readers Digest Birds of Canada. Edited by R. Greenberg 
et al. 2004. Dorling Kindersly India Ltd., c/o Tourmaline 
Editions, 662 King Street West, Suite 304, Toronto, Ontario 
MSV 1M7. 


Reptiles and Amphibians of the Amazon: An Ecotourist’s 
Guide. By R. D. Bartlett and P. Bartlett Florida University 
Presses, USA. 291 pages. $57 Canadian. Paper. 


Wild Down Under: The Natural History of Australasia. 
By Neil Nightingale, Jeni Clevers, Neil Pearson and Mary 
Summerill. BBC Video. 240 pages, $50 Canadian. Cloth. 


Botany 


The Wild Flowers of Britain and Ireland — The Complete 
Guide to the British and Irish Flora. By Marjorie Blamey, 
Richard Fitter and Alastair Fitter. A & C Black. 512 pages, 
$42 Canadian. Paper. 


Environment 


*Bull’s Eye — Unraveling the medical mystery of Lyme 
Disease. By Jonathon Edlow. 2004. Yale University Press, 
Box 209040, New Haven, Connecticut. 


+ Canoeing, Kayaking & Hiking Temagami. By Hap Wil- 
son. 2004. The Boston Mills Press (Firefly Books), 132 Main 
Street, Erin, Ontario. 112 pages, $24.95. 


+ Chicken Soup for the Fisherman’s Soul. By J. Canfield, 
M. Hansen, K. & Dahlynn McKowen. 2004. Health Com- 
munications Inc. Paper. 


2004 


Dictionary of Ecology. By Michael Allaby. 2004. Oxford 
University Press, 70 Wynford Drive, Don Mills, Ontario 
M3C 1J9. 448 pages, $29.95. 


+ The Importance of Species. Edited by P. Karliva and S. 
Levin. 2004. Princeton University Press, 41 William Street, 
Princeton, New Jersey. xvi + 427 pages, not illustrated, Paper. 


+ Lewis and Clarke on the Great Plains. By Paul Johns- 
gard. 2003. University of Nebraska Press. xiv + 143 pages, 
$14.95 U.S. 


+ Missinaibi. By Hap Wilson. 2004. The Boston Mills Press 
(Firefly Books), 132 Main Street, Erin, Ontario. 134 pages, 
$24.95. 


+ A place between the tides: A naturalist’s reflections on 
the salt marsh. By Harry Thurston. 2004. Greystone Books. 
Paper. $22.95 CAAD, $15.00 U.S. 


+ Politics of the Wild - Canada & Endangered Species. 
Edited by K. Beazley and R. Boardman. 2004. Oxford Uni- 
versity Press, 70 Wynford Drive, Don Mills, Ontario. x + 
254 pages, not illustrated, $29.95 Paper. 


Population Ecology: First Principles. By John H. Vander- 
meer and Deborah E. Goldberg. Princeton UP, USA. 280 
pages, $123 Cdn, Cloth. 


A Primer of Ecological Genetics. By J. Conner and D. Hartt. 
2004. Sinauer Associates, Box 407, Sunderland, Massachu- 
setts. 207 pages, $34.95 U.S., Paper. 


BOOK REVIEWS 


293 


* Resource and Environmental Management in Canada. 
Edited by B. Mitchell. 2004. Oxford University Press, 70 
Wynford Drive, Don Mills, Ontario. x + 600 pages, $29.95 
Paper. 


+ Rivers of the Upper Ottawa Valley. By Hap Wilson. 2004. 
The Boston Mills Press (Firefly Books), 132 Main Street, 
Erin, Ontario. 112 pages, $24.95. 


+ The Selbourne Pioneer. By T. Dodswell. 2004. Ashgate 
Publising Company, Suite 420, 101 Cherry St., Burlington, 
Vermont, USA. xvii + 238 pages, not illustrated. Paper. 


+ Shaped by the West Wind — Nature and History in 
Georgian Bay. By C. Campbell. 2004. UBC Press, 2029 
West Mall, Vancouver, British Columbia, V6T 172 Canada. 
256 pages, $85 Cdn. 


The Smithsonian Atlas of the Amazon. By Michael Gould- 
ing, Ronaldo Barthem and Efrem Ferreira. Smithsonian 
Books, Washington, D.C., USA. 255 pages, $75 Cdn. Cloth. 


+ Teaching in Eden. 2003. By J. Janovy Jr. Routledge, 29 
West 35th St., New York, New York 10001 USA. 187 pages, 
not illustrated, $22.95 U.S., Paper. 


+ Wilderness Rivers of Manitoba. By Hap Wilson and S. 
Aykroyd. 2004. The Boston Mills Press (Firefly Books), 
132 Main Street, Erin, Ontario, Canada. 112 pages, $24.95. 


+ Available 
* Assigned 


News and Comment 
Update on Quebec Amphibian and Reptile Atlas Project: Rana-Saura 6(2) 


The newsletter Rana-Saura resumes with Volume 6, Num- 
ber 2 issued for April 2004. The previous issue was published 
in March 2000. Due to translation expense no English ver- 
sion is currently available of the new issue (of 383 participants, 
only 23 are anglophones). Contents are: Et ¢a continue! — 
Atlas des amphibiens et des reptiles du Québec — Suivi des 
populations d’amphibiens du Québec — La présence de la Rai- 
nette faux-grillon boreale [Pseudarcris maculata] enfin con- 
firmée [Jean-Francois Desroches] — Un nouvelle reptile pour 
Québec! [Jean-Fran¢ois Desroches] — Le tout nouveau guide 
d’identification des amphibiens et des reptiles du Québec! 

Rana-Saura is edited by David Rodrigue, for the Atlas of 


amphibians and reptiles of Quebec (continuing in 2004 in its 
17" year; since 2002 the entries have increased from 22 000 
to nearly 45 000) and the Survey of populations of amphibians 
of Quebec (in 2004 there will be 45 participants conducting 
42 road surveys). These projects are coordinated by the Eco- 
musuem of the St. Lawrence Valley Natural History Society 
and financially aided by La Sociéte de la Faune et des Parcs 
du Québec. 

It is distributed by St. Lawrence Valley Natural History 
Society, 21-125 chemin Sainte-Marie, Sainte-Anne-de-Belle- 
vue, Quebec H9X 3Y7 Canada. Phone (514) 457-9449, 
extension 105: Fax (514) 457-0769; e-mail: ecomus @total.net. 


Froglog: Newsletter of the Declining Amphibian Populations Task Force (62) 


Number 62, April 2004. Contents: Southern African Frog 
Atlas Project completed! (Marius Burger and J. A. Harrison) 
— Mark Twain’s frog not croaked after all! (Robert Stack) — 
Decline of the Kihansi Spray Toad, Nectophrynoides asper- 
ginis from the Udzungwa Mountains, Tanzania (Che Weldon 
& Louis H. du Preez) — The Amphibian Fauna of Nagaland, 
India: Species and Habitats (Meren Ao & Sabitry Bordoloi) 
— Seed Grant News — Froglog Shorts. 

Froglog is the bi-monthly newsletter of the Declining Am- 
phibian Populations Task Force of The World Conservation 
Union (IUCN)/Species Survival Commission (SSC) and is 


supported by The Open University, The World Congress of 
Herpetology, and Arizona State University. The newsletter 
is edited by John W. Wilkinson, Department of Biological 
Sciences, The Open University, Walton Hall, Milton Keynes, 
MK7 6AA, United Kingdom; e-mail: daptf@open.ac.uk. 
Funding for Froglog is underwritten by the Detroit Zoo- 
logical Institute, P. O. Box 39, Royal Oak, Michigan 48068- 
0039, USA. Publication of issue 62 was also supported by 
Peace Frogs www.peacefrogs.com and by RANA and the U.S. 
National Science Foundation grant DEB-0130273. 


Amphipacifica: Journal of Aquatic Systematic Biology 3(4) 


Volume 3, Number 4, dated 30 March 2004, is 113 pages 
and contains the talitroidean amphipod family Najnidae in 
the North Pacific region: systematics and distributional ecol- 
ogy (E. L. Bousfield and Pierre Marcoux) and The amphi- 
pod family Pleustidae (mainly subfamilies Mesopleustinae. 
Neopleustinae, and Stenopleustinae) from the Pacific coast 
of North America: systematics and distributional ecology 


Marine Turtle Newsletter (104) 


April 2004. 36 pages: ARTICLES: Nesting of Hawksbill 
Turtles in Paraiba-Brazil: Avoiding Light Pollution Effects 
— Stranding of Small Juvenile Leatherback Turtle in West- 
ern Australia — Post-nesting Movements of Green Turtles 
Tagged in the Turtle Islands, Tawi-Tawi, Philippines — Notes 
from Preliminary Market Surveys in Morocco — NOTES: 
Recoveries of Two Post-hatching Loggerhead Turtles in the 
Northern Adriatic Sea — Green Turtle Nesting on the Gulf of 
Oman Coastline of the Islamic Republic of Iran — Confirmed 
Nesting of the Loggerhead Turtle in Corsica — Marine Tur- 
tles in Iran: Results from 2002 — Sightings of the Leatherback 
Turtle off the Southern Coast of Rio de Janeiro, Brazil — 


(E. A. Hendrycks and E. L. Bousfield). 

Amphipacifica (ISSN 1189-9905) is published quarterly by 
Amphipacifica Research Publications, Edward L. Bousfield, 
Managing Editor, 1710-1275 Richmond Road, Ottawa, On- 
tario K2B 8E3 Canada. Annual subscription rates are $40.00 
US or $50.00 Canadian funds. Back numbers are available 
at $10 each. 


MEETING REPORTS — IUCN/SSC MTSG News — BOOK REVIEWS 
— ANNOUNCEMENTS NEWS & LEGAL BRIEFS — RECENT PUBLI- 
CATIONS. 

The Marine Turtle Newsletter is edited by Brendan J. 
Godley and Annette C. Broderick, Marine Turtle Research 
Group, School of Biological Sciences, University of Exeter, 
Exeter EX4 4PS United Kingdom; e-mail MTN @seaturtle. 
org; Fax +44 1392 263700. Subscriptions and donations to- 
wards the production of both the MTN and its Spanish edi- 
tion NTM [Noticiero de Tortugas Marinas] should be made 
online at www.seaturtle.org/mtn or c/o SEATURTLE.ORG 
11400 Classical Lane, Silver Spring, Maryland 20901 USA. 


294 


2004 THE CANADIAN FIELD-NATURALIST 295 


Erratum 118(1): 80 


Sloan, N. A., and P. M. Garier. 2004. Introduced marine species in the Haida Gwaii (Queen Charlotte Islands) Region, British 
Columbia. Canadian Field-Naturalist 118(1): 77-84. 


Langara Island 
7 Parry Passage 
Cryptosula pallasiana He Rose 
Limnodrilus monothecus | Masset Spit 
/ Delkatla 


/ ; 
/ ® Pelonaia corrugata 


/ Inlet 
Heh Ve 


~ Trechus obtusus 


Limnodrilus monothecus 


om 


Cryptosula pallasiana . a a x 
\ 
xe Kumdis Bay 
\ Yakoun Bay 
Graham 


Rennell Sound o Island 
: Hecate 
Skidegate he 
Inlet 
A 


Moresby 


‘ Island 
Cartwright 
Sound 


Tasu 
Sound 


Bowerbankia gracilis 


Hotspring 
Ciona intestinalis Island 
Legend STUNG a 


a Sargassum muticum 
Dolomite "Burnaby" 
Narrows 


O Mya arenaria 
Qo oe 
A 
Harriet 


© Ampithoe vallida 
@ Other invertebrate sightings as noted B Harbour 
pl ceeeen eels 
at 
5a 
Bowerbankia gracilis ~~ — 


nt Houston Stewart 
Channel 


Schizoporella unicornis ® 


Tubularia crocea/ 
131°W 


FicureE |. Map of Haida Gwaii showing locations mentioned in the text and collection sites of introduced marine plant and 
invertebrate species, based on data from Sloan and Bartier (2000) and Sloan et al. (2001). 


Editor’s Report for Volume 117 (2003) 


Mailing dates for issues in volume 117 were: (1) 
30 September 2003, (2) 23 March 2004, (3) 12 July 
2004, (4) 16 September 2004. 

A summary of membership and subscriber totals 
2003 is given in Table 1. The number of articles and 
notes in volume 117 is summarized in Table 2 by topic; 
totals for Book Reviews and New Titles are given in 
Table 3, and the distribution of content by page totals 
per issue in Table 4. Council continued contribution of 
40% of membership dues for publication. All of sub- 
scriptions (both individual and institutional) also go 
toward publication. As well, the Council allocated 80% 
of the annual interest from the Manning Fund and 
other capital funds to The Canadian Field-Naturalist. 
The Manning fund portion is specifically to offset the 
publication cost of northern papers where author and 
institutional contributions were insufficient to cover 
page charges. 

Emil Holst arranged setting and printing at St. Joseph 
Print Group, Ottawa, for 117(1), with special thanks due 
Cameron Fraser and Willy Silfwerbrand, and at Gilmore 
Printers, Ottawa, for 117 (2,3,4) with special thanks to 
Wendy Cotie and Ally Reckzin. Leslie Cody prepared 
the Index, Elizabeth Morton read the galleys. Bill 
Cody as Business Manager handled all reprint requests 
and charges and oversaw and proofed the compilation 
of the Index. Wilson Eedy continued as Book-Review 
Editor until his death in June, and Roy John replaced 
him in August 2004. 

Manuscripts (excluding book reviews, notices, and 
reports) submitted to The Canadian Field-Naturalist 
totalled 84 in 2003, down 16 from 100 in 2002. The 
following reviewed for papers submitted in 2003: Asso- 
ciate Editors: (number of manuscripts reviewed in par- 
entheses): R. Anderson, Canadian Museum of Nature, 
Ottawa (5): C. D. Bird, Erskine, Alberta (14); B. W. 


TABLE 2. Number of articles and notes published in The Cana- 
dian Field-Naturalist Volume 117 (2003) by major field of 
study. 


Subject Articles Notes Total 
Mammals 19 * 14 33 
Birds Lie 6 23 
Amphibians + reptiles 4 1 BD) 
Fish 3 D. 5 
Invertebrates 6 1 6 
Plants** 16 3 19 
Tributes 2 0 2 
Totals 67 27 93 


*one article counted twice, one in each group: **includes one 
feature article. 


Coad, Canadian Museum of Nature, Ottawa (10); R. 
R. Campbell, St. Albert, Ontario (6); P. M. Catling, 
Agriculture and Agri-food Canada, Ottawa (11); A. J. 
Erskine, CWS, Sackville, New Brunswick (13); D. F. 
McAlpine, New Brunswick Museum, Saint John, N.B. 
(8); D. W. Nagorsen, Victoria, British Columbia (11); 
W. O. Pruitt, Jr., University of Manitoba, Winnipeg (21); 
Others: (one review each except as marked): P. Achuff, 
Waterton National Park, Alberta; Y. Alarie, Laurentian 
University, Sudbury, Ontario; N. Alfonso, Canadian 
Museum of Nature, Ottawa, Ontario; J. F Bain, Uni- 
versity of Lethbridge, Alberta; R. Bird, Cascadia Re- 
search, Olympia, Washington; K. Bellamy, OMNR, 
Peterborough; J. R. Bider, Ecomuseum, Ste-Anne-de- 
Bellevue, Quebec (3); E. L. Bousfield, Ottawa, On- 
tario; J. P. Bogart, University of Guelph, Ontario; I. 
Brodo, Canadian Museum of Nature, Ottawa, Ontario 
(2); R. J. Brooks, University of Guelph, Ontario; J. 
Calambokidis, Cascadia Research, Olympia, Washing- 
ton; S. E. Campana, Bedford Institute of Oceanography, 


TABLE |. The 2003 circulation of The Canadian Field-Naturalist (2002 in parenthesis). Totals compiled by W. J. Cody. Forty 
percent of membership dues and 100% of subscriptions go to publication of The Canadian Field-Naturalist. Members vote 


on Club affairs, subscribers and institutions do not. 


Canada USA Other Totals 
Memberships 
Family & individual E, (OMI) 36) G2) 7 (8) 838 (951) 
Subscriptions 
Individuals 163 (164) 60 (65) 6 (5) 229° (234) 
Institutions lesa — is(GuZAl)) 241 (252) 307762) 428 (455) 
Totals 1005 (1246) 337 (349) 43 (45) 1495 (1640) 


Note: 22 countries are included under “Other” (outside Canada and United States): Austria, Belgium, Brazil, Denmark (2), 
United Kingdom (9: including | to Scotland), Finland (2), France (3: including | to St. Pierre & Miquelon), Germany (2), 
Iceland, Ireland, Japan, Mexico, Netherlands (3), New Zealand, Norway (4), Poland, Russia, South Africa, Spain (3), 


Sweden (2), Switzerland (2), Trinidad and Tobago. 


296 


2003 


TABLE 3. Number of reviews and new titles published in Book 
Review section of The Canadian Field-Naturalist Volume 
117 by topic. 


Reviews New Titles 
Zoology 40 30 
Botany 11 23 
Environment 22 4] 
Miscellaneous 12 18 
Young Naturalists 0 18 
Totals 85 130 


Dartmouth, Nova Scotia; L. Carbyn, Canadian Wild- 
life Service, Edmonton, Alberta (4); A. Ceska, Victoria, 
British Columbia; A. H. Clarke, Jr., Portland, Texas; 
W. J. Cody, Agriculture and Agri-food Canada, Ottawa; 
V. Craig, EcoLogic Research, Gabriola Island, British 
Columbia; M. Crete, Ministere de 1’ Environnement et 
de la Faune, Quebec; E. J. Crossman, Royal Ontario 
Museum, Toronto, Ontario; S. J. Darbyshire, Agriculture 
and Agri-food Canada, Ottawa; R. Day, Canadian 
Museum of Nature, Ottawa; C. Ely, U. S. Fish and 
Wildlife Service, Anchorage, Alaska; M. Engstrom, 
Royal Ontario Museum, Toronto; B. Erickson, Salon. 
Ohio; M. Fairbarns, Conservation Data Centre, B. C. 
Government, Victoria; M. Gosselin, Canadian Museum 
of Nature, Ottawa; T. Goward, Clearwater, British 
Columbia; W. A. Fuller, Athabasca University, Alberta; 
D. Galbraith, Royal Botanical Gardens, Hamilton, 
Ontario; Marco Festa-Bianchet, University of Sher- 
brooke, Quebec; P. T. Gregory, University of Victoria, 
British Columbia (2); D. Gummer, Provincial Museum 
of Alberta, Edmonton (2); E. Haber, National Botani- 
cal Services, Ottawa; F. Harrington, Mt. St. Vincent 
University, Halifax, Nova Scotia; R. Harrington, Cana- 
dian Museum of Nature, Ottawa; D. Henry. Parks 
Canada, Haines Junction, Yukon Territory; S. Herrero, 
University of Calgary, Alberta; G. V. Hilderbrand, 
Alaska Department of Fish and Game, Anchorage; R. 
P. Hodge, Gig Harbor, Washington; C. S. Houston, 
Saskatoon, Saskatchewan (2); E. Holm, Royal Ontario 
Museum, Toronto; G. Jarrell, University of Alaska 
Museum, Fairbanks, Alaska; D. Johnson, Canadian 
Forest Service, Edmonton, Alberta; N. Jotham, NRJ 
Technical Services, Trade & Animal Welfare, Ottawa: 
R. B. King, Northern Illinois University, DeKalb, Illi- 
nois; J. L. Koprowski, Wildlife and Fisheries Resources, 
University of Arizona, Tucson; S. Lariviere, Delta 
Waterfowl Foundation, Portage La Prairie, Manitoba; 
D. Larson, Memorial University of Newfoundland, 
St. John’s; L. E. Licht, York University, North York, 
Ontario; J. Lien, Memorial University of Newfound- 
land, St. John’s; S. Lindgren, University of Northern 
British Columbia, Prince George; G. L. Mackie, Uni- 
versity of Guelph, Ontario; F. F. Mallory, Laurentian 
University, Sudbury, Ontario; A. Martel, Canadian 


EDITOR’S REPORT 


297 


TABLE 4. Number of pages per section published in The 
Canadian Field-Naturalist Volume 117 (2003) by issue. 


(1) (2) (3) (4) Total 
Articles WAI SB) 109 119 490 
Notes 12 16 15 17 60 
Tributes 1] 0 16 (0) Dil, 
Book Reviews’ 13 25 26 18 82 
CFN/OFNC Reports x 0 2 8 212. 
News and Comment 3 2 | 2 8 
Index 0) 0 0 30 30 
Advice to Contributors 0 0 | 0 | 
Totals 166 180 176 188 710 


*Total pages for book review section include both reviews 
and new titles listings. 

**Includes CFN Editors reports (issue 2), OFNC Annual 
Business Meeting (3) and OFNC Awards (4). 


Museum of Nature, Ottawa; John Maunder, New- 
foundland Museum, St. John’s, Newfoundland; M. K. 
McNicholl, Burnaby, British Columbia; G. R. Miche- 
ner, University of Lethbridge, Alberta; G. Mulligan, 
Agriculture and Agri-food Canada, Ottawa; E. Nol, 
Trent University, Peterbourgh; P. Paquet, Meacham, 
Saskatchewan; K. Prior, Canadian Wildlife Service, 
Ottawa; G. Proulx, Alpha Research & Management 
Ltd., Sherwood Park, Alberta; C. Renaud, Canadian 
Museum of Nature, Ottawa, Ontario; B. Riddell, Paci- 
fic Biological Station, Nainimo, British Columbia; R. 
A. Ring, University of Victoria, British Columbia; 
James Schaefer, Trent University. Peterborough, On- 
tario; W. B. Schofield, University of British Columbia, 
Vancouver; F. W. Schueler, Oxford Station, Ontario; 
B. Slough, Whitehorse, Yukon Territory; I. Stirling, 
Canadian Wildlife Service, Edmonton, Alberta (2); J. 
Straley, University of Alaska Southeast, Sitka, Alaska; 
D. Strickland, Dwight, Ontario; E. S. Telfer, Edmon- 
ton, Alberta; I. Thompson, Canadian Forest Service, 
Sault Ste. Marie, Ontario; C. Traynor, Ottawa, Ontario; 
N. Vachon, Sociéte de la faune et des parcs du Québec, 
Longueuil; P. Weigl, Wake Forest University, Winston- 
Salem, North Carolina; G. Wobeser, Western College 
Veterinary Medicine, University of Saskatchewan, Sas- 
katoon; J. N. Womble, University of Alaska-Fairbanks, 
Juneau, Alaska; P. M. Youngman, Ottawa, Ontario (2). 

I am also indebted to the President of the Ottawa 
Field-Naturalists’ Club Gary McNulty and the Club 
Council for continuing support of the journal; Chair- 
man Ron Bedford and the Publications Committee of 
the OFNC for editorial encouragement and support, to 
the Canadian Museum of Nature for access to its library 
and the facilities at the Natural Heritage Building, 
1740 Pink Road, Aylmer, Quebec, and to Joyce for 
everything else. 


FRANCIS R. COOK 
Editor 


Advice for Contributors to The Canadian Field-Naturalist 


Content 

The Canadian Field-Naturalist is a medium for the publi- 
cation of scientific papers by amateur and professional natu- 
ralists or field-biologists reporting observations and results 
of investigations in any field of natural history provided that 
they are original, significant, and relevant to Canada. All read- 
ers and other potential contributors are invited to submit for 
consideration their manuscripts meeting these criteria. The 
journal also publishes natural history news and comment items 
if judged by the Editor to be of interest to readers and sub- 
scribers, and book reviews. Please correspond with the Book 
Review Editor concerning suitability of manuscripts for this 
section. For further information consult: A Publication Policy 
for the Ottawa Field-Naturalists’ Club, 1983. The Canadian 
Field-Naturalist 97(2): 231-234. Potential contributors who 
are neither members of The Ottawa Field-Naturalists’ Club 
nor subscribers to The Canadian Field-Naturalist are encour- 
aged to support the journal by becoming either members or 
subscribers. 


Manuscripts 

Please submit, to the Editor, in either English or French, 
three complete manuscripts written in the journal style. 
The research reported should be original. It is recommended 
that authors ask qualified persons to appraise the paper before 
it is submitted. All authors should have read and approved it. 
Institutional or contract approval for the publication of the data 
must have been obtained by the authors. Also authors are ex- 
pected to have complied with all pertinent legislation regard- 
ing the study, disturbance, or collection of animals, plants or 
minerals. The place where voucher specimens have been de- 
posited, and their catalogue numbers, should be given. Lati- 
tude and longitude should be included for all individual local- 
ities where collections or observations have been made. 

Print the manuscript on standard-size paper, doublespace 
throughout, leave generous margins to allow for copy mark- 
ing, and number each page. For Articles and Notes provide 
a citation bibliographic strip, an abstract and a list of key 
words. Generally, words should not be abbreviated but use 
SI symbols for units of measure. The names of authors of 
scientific names may be omitted except in taxonomic 
manuscripts or other papers involving nomenclatural 
problems. “Standard” common names (with initial letters 
capitalized) should be used at least once for all species of 
higher animals and plants; all should also be identified by 
scientific name. 

The names of journals in the Literature Cited should be 
written out in full. Unpublished reports and web documents 
should not be cited here but placed in the text or in a sepa- 
rate Documents Cited section. List the captions for figures 
numbered in arabic numerals and typed together on a separate 
page. Present the tables each titled, numbered consecutively 
in arabic numerals, and placed on a separate page. Mark in 
the margin of the text the places for the figures and tables. 


Check recent issues (particularly Literature Cited) for 
journal format. Either “British” or “American” spellings are 
acceptable in English but should be consistent within one 
manuscript. The Oxford English Dictionary, Webster’s 
New International Dictionary and le Grand Larousse 
Encyclopédique are the authorities for spelling. 


Illustrations 

Photographs should have a glossy finish and show sharp 
contrasts. Electronic versions should be high resolution. Photo- 
graphic reproduction of line drawings, no larger than a 
standard page, are preferable to large originals. Prepare line 
drawings with India ink on good quality paper and letter (don’t 
type) descriptive matter. Write author’s name, title of paper, and 
figure number on the lower left corner or on the back of each 
illustration. 


Reviewing Policy 

Manuscripts submitted to The Canadian Field-Naturalist 
are normally sent for evaluation to an Associate Editor (who 
reviews it or asks another qualified person to do so), and at 
least one other reviewer, who is a specialist in the field, cho- 
sen by the Editor. Authors are encouraged to suggest names 
of suitable referees. Reviewers are asked to give a general 
appraisal of the manuscript followed by specific comments 
and constructive recommendations. Almost all manuscripts 
accepted for publication have undergone revision—sometimes 
extensive revision and reappraisal. The Editor makes the 
final decision on whether a manuscript is acceptable for pub- 
lication, and in so doing aims to maintain the scientific quality, 
content, overall high standards and consistency of style, of 
the joumal. 


Special Charges — Please take note 

Authors must share in the cost of publication by paying 
$80 for each page, plus $15 for each illustration (any size up 
to a full page), and up to $80 per page for tables (depending 
on size). Authors may also be charged for their changes in 
proofs. Reproduction of color photos is extremely expensive; 
price quotations may be obtained from the Business Manager. 
If grant or institutional funds are not available, club members 
and subscribers may apply for a waiver of charges for the 
first five pages. 

Limited joumal funds are available to help offset publi- 
cation charges to authors with minimal financial resources. 
Requests for financial assistance should be made to the Busi- 
ness Manager when the manuscript is accepted. 


Reprints 
An order form for the purchase of repents will accompany 
the galley proofs sent to the authors. 


FRANCIS R. CooK, Editor | 
RR 3 North Augusta, Ontario KOG TRO Canada | 


298 


TABLE OF CONTENTS (concluded) Volume 118 Number 2 


Notes 


Observations of-above surface littoral foraging in two sea ducks, Barrow’s Goldeneye, 
Bucephala islandica, and Surf Scoter, Melanitta perspicillata, in coastal southwestern 
British Columbia DEBORAH L. LACROIX, KENNETH G. WRIGHT, and DANIEL KEN1 


New records of Cyperaceae and Juncaceae from the Yukon Territory 
STUART A. HARRIS and PETER W. BALL 


“Ashkui”, vernal ice-cover phenomena and their ecological role in southern Labrador 
SHAUNA M. BAILLIE, CORINNE D. WILKERSON, and TINA L. NEWBURY 


Sea Otter, Enydra lutris, sightings off Haida Gwaii/Queen Charlotte Islands, Canada, 1972-2002 
KIMBERLY RAUM-SURYAN, KENNETH PITCHER, and RICHARD LAMY 


iFirst record of a Barred Owl, Strix varia, in Labrador ISABELLE SCHMELZER and FRANK PHILLIPS 


Forked Three-awned Grass, Aristida basiramea Engelm. ex Vasey: a new addition to the 
flora of Quebec JACQUES BRISSON 


Evidence for the use of vocalization to coordinate the killing of a White-tailed Deer, Odocoileus 
virginianus, by Coyotes, Canis latrans ERICH M. MUNTZ and BRENT R. PATTERSON 


Book Reviews 

ZOoLoGy: Fish of Alberta — Sharks — In Search of the Golden Frog — Conversations with an Eagle — 
North American Owls, Biology and Natural History, Second Edition — The Mountain White-Crowned 
Sparrow: Migration and Reproduction at High Altitude — Seabird Bycatch: Trends, Roadblocks, and 
Solutions — The Life of Mammals — Geographic Variation in Size and Shape of Savannah Sparrows 
(Passerculus sandwichensis) — Warblers of the Great Lakes and Eastern North America — Sparrows 
and Finches of the Great Lakes and Eastern North America — Birds of the Yukon Territory 


Botany: Wild Flowers of the Yukon, Alaska & Northwestern Canada, 2™ edition — Wild Flowers of Field 
| & Slope in the Pacific Northwest — Wild Flowers of Forest & Woodland in the Pacific Northwest — 
Wild Flowers of the Mountains in the Pacific Northwest 


MISCELLANEOUS: The Canoe: A Living Tradition — A Passion for Wildlife: The History of the Canadian 
Wildlife Service — Birding on Borrowed Time 


New TITLES 


News and Comment 

Update on Quebec Amphibian and Reptile Atlas Project: Rana-Saura 6(2) — Froglog: Newsletter of the 
Declining Amphibian Populations Task Force (62) — Amphipacifica: Journal of Aquatic Systematic 
Biology 3(4) — Marine Turtle Newsletter (104) — Erratum: Sloan and Bartiers 118(1): 80 


| Editor’s Report Volume 117 (2003) FRANCIS R. COOK 


| Advice for Contributors 


Mailing date of the previous issue 118(1): 15 December 2004 


2004 


264 


266 


THE CANADIAN FIELD-NATURALIST Volume 118 Number 2 2004 


Articles 


Morphology and population characteristics of Vancouver Island Cougars, 
Puma concolor vancouverensis STEVEN F. WILSON, APRYL HAHN, AARON GLADDERS, | 
KAREN M. L. Gou, and Davip M. SHACKLETON 159 


Conservation evaluation of Lemmon’s Holy Fern, Polystichum lemmonii, a threatened 
fern in Canada GEORGE W. DOUGLAS 164 


Conservation evaluation of the Pacific population of Tall Woolly-heads, Psilocarphus elatior, 
an endangered herb in Canada GEORGE W. DOUGLAS and JEANNE M. ILLINGWORTH 169 


Conservation evaluation of Howell’s Triteleia, Triteleia howellii, an endangered lily 
in Canada GEORGE W. DOUGLAS and JENIFER L. PENNY 174 


Conservation evaluation of Small-flowered Lipocarpha, Lipocarpha micrantha (Cyperaceae), 
in Canada TYLER W. SMITH, GEORGE W. DOUGLAS, and ALLAN G. HARRIS 175) 


Immobilizatin of clover-trapped White-tailed Deer, Odocoileus virginianus, with medetomidine 
and ketamine, and antagonism with atipamezole JOSHUA J. MILLSPAUGH, 
BRIAN E. WASHBURN, TAMARA M. MEYER, JEFF BERINGER, and LONNIE P. HANSEN 185 


Predation on two Mule Deer, Odocoileus hemionus, by a Canada Lynx, Lynx canadensis, in the southern 
Canadian Rocky Mountains DorTE PosziG, CLAYTON D. Apps, and ALAN DIBB 191 


Effect of fire intensity and depth of burn on Lowbush Blueberry, Vaccinium angustifolium, and 
Velvet Leaf Blueberry, Vaccinium myrtilloides, production in eastern Ontario 
Luc C. DUCHESNE and SUSANNE WETZEL 195 


Spring dispersal patterns of Red-winged Blackbirds, Agelaius phoeniceus, staging in eastern 
South Dakota H. J. HomMaAN, G. M. Linz, R. M. ENGEMAN, and L. B. PENRY 201 1f 


Changes in loon (Gavia spp.) and Red-necked Grebe (Podiceps grisegena) populations in the Lower 
Matanuska-Susita Valley, Alaska TAMARA K. MILLS and BRAD A. ANDRES 210 


The Barred Owl, Strix varia, in Alberta: distribution and status LisA TAKATS PRIESTLEY 215 


Use of Eelgrass, Zostera marina, wrack by three species of ladybird beetles (Coleoptera: 
Coccinellidae) in Prince Edward Island DaviD J. GARBARY, SARAH FRASER, 
CARRIE FERGUSON, and RANDOLPH F. LAUFF 225 


Predicting the effects of Cerulean Warbler, Dendroica cerulea, management on eastern Ontario 
bird species JASON JONES, WILLIAM J. MCLEISH, and RALEIGH J. ROBERTSON Ds 


Parasite prevalence in Dark-eyed Juncos, Junco hyemalis, breeding at different elevations i 
HEATHER BEARS 235 iF 


Movements of subadult male Grizzly Bears, Ursus arctos, in the central Canadian Arctic 
ROBERT J. GAU, PHILIP D. MCLOUGHLIN, RAY CASE, H. DEAN CLUFF, 
ROBERT MULDERS, and FRANCOIS MESSIER 239 


Severe chronic neck injury caused by a snare in a Coyote, Canis latrans 
PIERRE- Y VES DaAousT and PETER H. NICHOLSON 243 


Interactions of Brown Bears, Ursus arctos, and Gray Wolves, Canis lupus, at Katmai National Park and 
Preserve, Alaska Tom S. SMITH, STEVEN T. PARTRIDGE, and JOHN W. SCHOEN 247 


Wild Turkey, Meleagris gallopavo silvestris, behavior in central Ontario during winter 
LINH P. NGUYEN, JOSEPH HAMR, and GLENN H. PARKER 25 


(continued on inside back cover, 


ISSN 0008-3550 


| The CANADIAN 
—  FIELD-NATURALIST 


Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada 


Volume 118, Number 3 July-September 2004 


The Ottawa Field-Naturalists’ Club 


FOUNDED IN 1879 


Patrons 
Her Excellency The Right Honourable Adrienne Clarkson, C.C., C.M.M., C.D. 
Governor General of Canada 
His Excellency John Ralston Saul, C.C. 


The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to 
encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields 
as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environments 
of high quality for living things. 


Honorary Members 


Edward L. Bousfield Bruce Di Labio John A. Livingston E. Franklin Pope 
Donald M. Britton R. Yorke Edwards Stewart D. MacDonald William O. Pruitt, Jr. 
Irwin M. Brodo Anthony J. Erskine Hue N. MacKenzie Joyce and Allan Reddoch 
William J. Cody John M. Gillett Theodore Mosquin Mary E. Stuart 
Francis R. Cook C. Stuart Houston Eugene G. Munroe John B. Theberge 
Ellaine Dickson George F. Ledingham Robert W. Nero Sheila Thomson 
2004 Council 

President: Mike Murphy Ronald E. Bedford Barbara Gaertner Stanley Rosenbaum 
Vice-President: Gillian Marston Fenja Brodo Diane Lepage Louise Schwartz 
Recording Secretary: Susan Laurie-Bourque John Cameron Christina Lewis David Smythe 
Treasurer: Frank Pope William J. Cody Karen McLachalan Hamilton Henry Steger 
Past President: Gary McNulty Kathy Conlan David Hobden Chris Traynor 

Francis R. Cook Cendrine Huemer 


To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, P.O. Box 35069, Westgate 
P.O. Ottawa, Canada K1Z 1A2, or e-mail: ofnc @achilles.net. 
For information on Club activities telephone (613) 722-3050 or check www.ofnc.ca 


The Canadian Field-Naturalist 


The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this 
journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. 


We acknowledge the financial support of the Government of Canada toward our mailing cost through the Publication Assistance 
Program (PAP), Heritage number 09477. 


Editor: Dr. Francis R. Cook, R.R. 3, North Augusta, Ontario KOG IRO; (613) 269-3211; e-mail: feook @achilles.net 
Copy Editor: Elizabeth Morton 

Business Manager: William J. Cody, P.O. Box 35069, Westgate P.O. Ottawa, Canada KIZ 1A2; (613) 759-1374 
Book Review Editor: Roy John, 2193 Emard Crescent, Ottawa, Ontario K1J 6K5, e-mail: roy.john@pwegsc.ge.ca 


Associate Editors: Robert R. Anderson Paul M. Catling David Nagorsen 
Charles D. Bird Brian W. Coad Donald F. McAlpine 
Robert R. Campbell Anthony J. Erskine William O. Pruitt, Jr. 


Chairman, Publications Committee: Ronald E. Bedford 


All manuscripts intended for publication except Book Reviews should be addressed to the Editor and sent by postal 
mail. Book-review correspondence should be sent by e-mail or postal mail to Roy John, Book-review Editor. 


Subscriptions and Membership 

Subscription rates for individuals are $28 per calendar year. Libraries and other institutions may subscribe at the rate of $45 per 
year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $28 (individual) $30 (family) $50 (sustaining) and 
$500 (life) includes a subscription to The Canadian Field-Naturalist. All foreign subscribers and members (including USA) must 
add an additional $5.00 to cover postage. The club regional journal, Trail & Landscape, covers the Ottawa District and Local Club 
events. It is mailed to Ottawa area members, and available to those outside Ottawa on request. It is available to Libraries at $28 per 
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Cover: International zone on rugged rock headland of the Queen Charlotte Islands, British Columbia, each dominant species cre- 
ating a distinct zone. The white lichen is mainly Coccotrema maritimum and the black one is almost entirely Verru cana 
maura. Below the Verru cana is a zone bare of both lichens and marine algae, and, still lower, the algae take over. Figure 
64 (page 77). Lichens of North America. By I. M. Brodo, S. D. Sharnoff, and S. Sharnoff. 2001. Yale University Press, New 
Haven, Connecticutt. See article by Brodo and Sloan on lichen zonatum pages 405-424. 


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| Volume 118, Number 3 


The Canadian Field-Naturalist vA. 


July—September 2004 


Population Dynamics of Deer Mice, Peromyscus maniculatus, and 
Yellow-pine Chipmunks, Tamias amoenus, in Old Field and Orchard 
Habitats 


THOMAS P. SULLIVAN!, DRUSCILLA S. SULLIVAN?, and EUGENE J. HOGUE? 


'Agroecology Program, Faculty of Land and Food Systems, University of British Columbia, Vancouver, British Columbia 
V6T 1Z4 Canada (corresponding author). 

? Applied Mammal Research Institute, 11010 Mitchell Avenue, Summerland, British Columbia VOH 1Z8 Canada 

3 Pacific Agri-Food Research Centre, Agriculture and Agri-Food Canada, 4200 Highway 97, Summerland, British Columbia 
V1X 7S3 Canada 


Sullivan, Thomas P., Druscilla S. Sullivan, and Eugene J. Hogue. 2004. Population dynamics of Deer Mice, Peromyscus mani- 
culatus, and Yellow-pine Chipmunks, Tamias amoenus, in old field and orchard habitats. Canadian Field-Naturalist 
118(3): 299-308. 


There are often several rodent species included in the small mammal communities in orchard agro-ecosystems. This study was 
designed to test the hypothesis that the population levels of Deer Mice (Peromyscus maniculatus) and Yellow-pine Chipmunks 
(Tamias amoenus) would be enhanced in old field compared with orchard habitats. Rodent populations were intensively live- 
trapped in replicate old field and orchard sites over a four-year period at Summerland, British Columbia, Canada. Deer Mouse 
populations were, on average, significantly higher (2.5 — 3.4 times) in the old field than orchard sites in summer and winter 
periods. Mean numbers/ha of Deer Mice ranged from 12.1 to 60.4 in old field sites and from 3.3 to 19.9 in orchard sites. 
Breeding seasons in orchards were significantly longer than those in old field sites, in terms of proportion of reproductive 
male Deer Mice. Recruitment of new animals and early juvenile survival of Deer Mice were similar in orchard and old field 
sites. Populations of Yellow-pine Chipmunks ranged in mean abundance/ha from 5.6 — 19.0 in old field sites and from 1.9 — 
17.5 on one orchard site, with no difference in mean abundance in 2 of 4 years of the study. Recruitment and mean survival 
of Yellow-pine Chipmunks also followed this pattern. This study is the first detailed comparison of the population dynamics 
of these rodent species in old field and orchard habitats. These species should be able to maintain their population levels and 
help contribute to a diversity of small mammals in this agrarian landscape. 


Key Words: Deer Mouse, Peromyscus maniculatus, Yellow-pine Chipmunk, Tamias amoenus, abundance, demography, old 


field, orchard, population dynamics, recruitment, reproduction, British Columbia. 


Voles of the genus Microtus are often the major 
rodent species in perennial grasslands and agro-eco- 
systems. In addition to grasses and herbs, these micro- 
tines feed on vascular tissues of tree and other crop 
plants primarily during winter months (Byers 1985; 
Lewis and O’Brien 1990). Other major rodent species 
include the Deer Mouse (Peromyscus maniculatus) 
and Yellow-pine Chipmunk (TZamias amoenus), which 
may occur in tree fruit orchards and adjacent habitats 
in the inland regions of the Pacific Northwest of 
North America. Neither of these associated species is 
known to damage orchard trees by their feeding activity 
as they are granivorous or insectivorous in their feeding 
habits (Baker 1968; Webster and Jones 1982; Sutton 
1992). 

Deer Mice or White-footed Mice (P. leucopus) com- 
monly occur with Microtus in old fields and other 
perennial grassland habitats (Tamarin 1977; Krebs 
1979; Dueser et al. 1981; Sullivan and Krebs 1981; 


Schweiger et al. 2000; Manson et al. 2001; Pearson et 
al. 2001) and in orchard agro-ecosystems (Sullivan et 
al. 1998). Few studies of voles in orchards have report- 
ed on the population dynamics of other species in the 
small mammal community. Sullivan et al. (1998) dis- 
cussed the population responses of all species to vege- 
tation management in apple (Malus domestica) orchards 
and Sullivan et al. (2000) reported changes in species 
diversity of these communities in orchards and old fields. 

Orchard habitats usually have frequent mowing and 
vegetation management treatments during summer 
months, and hence conditions for small mammals are 
more changeable than in old fields. Thus, this study 
was designed to test the hypothesis that the popula- 
tion dynamics of Deer Mice and Yellow-pine Chip- 
munks would be enhanced in old field compared with 
orchard habitats. To evaluate this hypothesis, we pro- 
vide a detailed analysis of the population dynamics of 
these species in the two habitats. 


299 


300 


Materials and Methods 
Study area and experimental design 

This study was conducted at the Pacific Agri-Food 
Research Centre in the Okanagan Valley, Summerland, 
British Columbia, Canada. The experimental design 
consisted of two replicate “old field” and two repli- 
cate orchard habitats. The old field habitats were 
abandoned (225 years) hay fields composed of Crest- 
ed Wheatgrass (Agropyron cristatum), Quack Grass 
(Agropyron repens), Downy Brome (Bromus tecto- 
rum), Diffuse Knapweed (Centaurea diffusa), with 
some minor herbaceous species such as Yellow Salsify 
(Tragopogon dubius), Great Mullein (Verbascum 
thapsus), American Vetch (Vicia americana), Prickly 
Lettuce (Lactuca serriola), and Tall Tumble-mustard 
(Sisymbrium altissimum). These old field sites were 
each 2 to 3 ha in area within a mosaic of sagebrush 
(Artemisia tridentata), Ponderosa Pine (Pinus pon- 
derosa) forest, and orchard habitats. These old fields 
had resident populations of Deer Mice and Yellow- 
pine Chipmunks. Other species included the Montane 
Vole (Microtus montanus), Great Basin Pocket Mouse 
(Perognathus parvus), Western Harvest Mouse (Reith- 
rodontomys megalotis), and a few Long-tailed Voles 
(M. longicaudus). 

The orchards were: (A) a five-year-old apple orchard 
unit, and (B) a 10-year-old apple orchard combined 
with a 15-year-old pear (Pyrus sp.) orchard as one 
unit. Both 1.2-ha orchards were located within a 90- 
ha mosaic of tree fruits and vineyards. Thus, our ex- 
perimental design had two true replicates of old field 
sites and two replicates of orchard sites. Each pair of 
old field and orchard sites was spatially segregated to 
enhance statistical independence (Hurlbert 1984). A 
third replicate pair would have strengthened the study 
but was not possible within the operational setting of 
the Research Centre. 

Common grass species on the orchard sites included 
Orchard Grass (Dactylis glomerata), Quack Grass, 
bluegrass (Poa spp.), Smooth Brome (Bromus inermis), 
and Crested Wheatgrass. These orchards were mowed 
five or six times each summer. Rodenticides were 
applied 3-4 times each winter in poison-bait feeder 
stations (Radvanyi 1974) for voles (Mouse Bait IT® — 
zinc phosphide and Ramik Brown® — diphacinone). 
Rodenticides were not present in the old field sites. 


Deer Mouse and Yellow-pine Chipmunk populations 
All animals were live-trapped on 1-ha grids with 
49 (7 x 7) trap stations located at 14.3-m intervals 
with one or two Longworth live-traps at each station. 
One of the old-field grids was an irregular shaped 
rectangle of 1 ha with the same 49 stations. The four 
grids (2 orchard and 2 old field) were live-trapped at 
3-week (spring, summer, and fall) and at 4- to 6-week 
(winter) intervals from June 1982 to April 1986. 
Traps were baited with whole oats, peanut butter, 
and carrot; coarse brown cotton was supplied as bed- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


ding. Traps were set on day 1, checked on the morning 
and afternoon of day 2 and morning of day 3, and then 
locked open between trapping periods. All animals 
captured were ear-tagged with serially numbered tags, 
breeding condition noted, weighed on Pesola spring 
balances, and point of capture recorded. The duration 
of the breeding season was noted by palpation of male 
testes and the condition of mammaries of the females 
(Krebs et al. 1969). A pregnancy was considered suc- 
cessful if a female was lactating during the period 
following the estimated time of birth of a litter. Ani- 
mals were released on the grids immediately after 
processing. 

Seasons were defined as summer (April to Septem- 
ber) and winter (October to March) periods. Thus, 
there were four summer and four winter periods from 
1982 to 1986. We used mass at sexual maturity to 
infer age classes of animals. Body mass was used as 
an index of age. The percentage of sexually mature 
animals was used to determine the mass limitations 
for juveniles and adults assuming that juveniles were 
seldom, if ever, sexually mature, and that at least 50% 
of the adults were sexually mature in their lowest 
mass class. Deer Mice (juvenile = 1 — 20 g, adult > 21 g) 
and Yellow-pine Chipmunks (juvenile = 1 — 44 g, adult 
2 45 g) were classified as juvenile or adult by body 
mass. Juveniles were considered to be young animals 
recruited during the study. Recruits were defined as 
new animals that entered the population through repro- 
duction and immigration. All handling of animals was 
in accordance with the principles of the Animal Care 
Committee, University of British Columbia. 


Demographic parameters 

Population densities were estimated by the Jolly- 
Seber model for reasons indicated by Jolly and Dick- 
son (1983). The Jolly-Seber (J-S) model provides the 
best.estimates of population size for mark and recap- 
ture data when trappability values are generally < 70% 
(Hilborn et al. 1976). However, when population size 
falls very low and no marked animals are recaptured, 
the J-S estimate becomes unreliable and impossible to 
calculate (Krebs et al. 1986). For these sample weeks, 
a minimum number of animals known to be alive 
(MNA) (Krebs 1966) value was substituted for a bio- 
logically unreasonable J-S estimate. 

Measurements of recruitment, number of lactating 
females, and early juvenile survival were derived from 
the sample of animals captured in each trapping ses- 
sion and then summed for summer periods. Early juve- 
nile survival is an index relating recruitment of young 
into the trappable population to the number of lactat- 
ing females (Krebs 1966). A modified version of this 
index is number of juvenile animals at week ft divided 
by the number of lactating females caught in week 
t — 3. Mean survival rates (28-day) for summer and 
winter periods were estimated from the Jolly-Seber 
model. Mean body mass of combined males and females 


2004 


SULLIVAN, SULLIVAN, AND HOGUE: POPULATION DYNAMICS OF DEER MICE AND CHIPMUNKS 30] 


Peromyscus maniculatus 


Number of animals/na 
NO (Se) A on oO) 
(eo) (ep) (ep) (oe) (=) 


= 
(eo) 


--Old field 1 Old field 2 
Orchard 1 Orchard 2 


J SNJIMMJSNJIMMJISNJIMM JSNJIMM 


1982 1983 


1984 1985 1986 


FiGurRE |. Population densities (Jolly-Seber) per ha of Deer Mice in replicate old field and orchard sites during the study. 
Shaded bars indicate winter periods. Months of year are represented by J = July; S = September; N = November; J 


January; M = March; M = May. 


was used as an index of condition within populations 
of Deer Mice and Yellow-pine Chipmunks during sum- 
mer and winter periods. 


Statistical analysis 

Mean trappability, mean abundance, mean number 
of recruits, mean Jolly-Seber survival rates, and mean 
body mass were evaluated by 95% confidence inter- 
vals (CI) for Deer Mice and Yellow-pine Chipmunks 
in old field and orchard sites during summer and winter 
periods, as per the recommendations of Gerard et al. 
(1998) and Johnson (1999). Proportion of adult males 
and adult females breeding was analyzed by a Chi- 
square 2 x 2 contingency table (Zar 1984) for each of 
the four summer periods. These datasets of proportion 
of animals breeding often include animals captured 
more than once, and hence they are not completely 
independent. Thus, the Chi-square analyses provide 
only an indication of the degree of difference between 
datasets. In all analyses, the level of significance was 
E1053 


Results 
Deer Mouse populations 

Totals of 751 and 459 individual Deer Mice were 
captured on the two old field and two orchard sites, 
respectively. Mean J-S trappability estimates for Deer 
Mice ranged from 62.6% to 73.8% in summer and 
from 46.6% to 76.6% in winter in the old field sites. 
These estimates in the orchard sites ranged from 48.0% 
to 81.7% in summer and from 39.0% to 80.8% in 
winter. 

Deer Mouse populations were, on average, general- 
ly higher (2.5 — 3.4 times) in the old field than orchard 
sites in summer and winter periods (Figure 1). There 
were some exceptions to this pattern in summers 1982 
and 1985 and winter 1985-1986 when mean abun- 
dance of Deer Mice was similar, at least in one old 
field—orchard site comparison (Table 1). Numbers 
of Deer Mice reached annual autumn peaks ranging 
from 30.8 to 69.3 animals/ha in the old field-1 site and 
from 57.2 to 67.5 animals/ha in the old field-2 site. 


302 THE CANADIAN FIELD-NATURALIST Vol. 118 


TABLE |. Mean abundance per ha + 95% confidence intervals for Deer Mice during summer and winter periods in replicate 
old field and orchard sites. Sample size (n = number of trapping periods) in parentheses. 


Old field Orchard 
Period 1 2 1 2 
Summer 1982 ye LES BikD 15.2 12.6 
(7) 95% CI 11.2 — 23.8 22, 7.7 —22.7 9.3 — 16.0 
Winter 1982-83 NG 25.8 49.0 16.8 6.0 
(5) 95% CI 21.9 — 29.7 43.0 —55.] 10.6 — 23.0 2.6 —9.4 
Summer 1983 Ve 41.9 55.6 15:5 19.9 
(9) 95% CI 36.2 — 47.6 50.1 — 61.2 8.3 — 22.8 14.4 — 25.4 
Winter 1983-84 xr 45.7 B92 10.6 9.0 
(8) 95% CI 39.8 — 51.6 29.5 — 48.8 8.7 — 12.5 2.3 — 15.7 
Summer 1984 1 ba 60.4 33.8 19.7 aL 
(8) 95% Cl 54.5 — 66.3 24.0 — 43.6 16.1 — 23.3 14.3 —21.0 
Winter 1984-85 x2 34.3 SES ite 333 
(6) 95% CI 22.4 — 46.1 5.1-21.8 4.7-17.5 2.2-4.4 
Summer 1985 V2 36.3 12.1 125 6.9 
(9) 95% Cl 31.9 — 40.7 7.9 — 16.2 5.4 — 19.6 3.7— 10.1 
Winter 1985-86 yf 41.1 22:2 19.6 4.4 
(7) 95% CI 36.9 — 45.3 6.2 — 38.2 10.5 — 28.7 1.8—7.0 
These old field populations also had increased recruit- In terms of mean abundance, Deer Mouse numbers 


ment in early summer (April — May) in some years _ in old field sites ranged from a low of 12.1/ha to a high 
which was likely related to the decline in breeding — of 60.4/ha during the four-year study (Table 1). Mean 
activity by mid-summer (see Figure 2). abundance of Deer Mice on the two orchard sites 


Peromyscus maniculatus 
Old field 1 +2 


80 


fe)) 
oO 


& 
jo) 


NO 
fo) 


Percentage of adult males breeding 


! 


JSNJIMMJISNJIMMJISNJIMMJISNJIMM 
1982 1984 1985 1986 


FiGurE 2. Percentage of adult males in reproductive condition for Deer Mice for pooled data in old field and orchard sites 
during breeding seasons each year. Shaded bars indicate winter periods. Months of year are represented by J = July: 
S = September; N = November; J = January; M = March; M = May. 


2004 


ranged from a low density of 3.3/ha to a high of 19.9/ha. 
Orchard Deer Mice had annual increases in recruitment 
in early summer with generally higher populations 
(1.5 times) in summer than winter (Figure 1, Table 1). 


Demographic parameters 

Adult male Deer Mice commenced breeding in 
January-February each year and continued up to July 
in the old field sites and through the summer and 
autumn months in the orchard sites (Figure 2). This 
longer breeding season in orchard than old field sites 
was significantly different in terms of proportion of 
reproductive males in 3 of 4 years (Table 2). Propor- 
tion of breeding female Deer Mice tended to follow 
this pattern but there was a significant difference 
between sites in 1984 only (Table 2). 

The pattern of recruitment in terms of mean num- 
bers of new Deer Mice was generally similar in old 
field and orchard sites in both summer and winter 
periods (Table 3). There was only one period (summer 


SULLIVAN, SULLIVAN, AND HOGUE: POPULATION DYNAMICS OF DEER MICE AND CHIPMUNKS 


303 


1985) when an old field — orchard comparison of mean 
values did not have overlapping 95% CI (Table 3). 
Thus, productivity within orchard populations of Deer 
Mice appeared to approach that of old field popula- 
tions, at least with respect to breeding potential and 
production of young. 

Over the four breeding seasons, early juvenile sur- 
vival was similar with overall mean values of 3.64 and 
2.12 young Deer Mice captured per lactating female 
in old field and orchard sites, respectively. Mean total 
survival of Deer Mice was consistently higher in the 
two old field sites and orchard-1 site than in the orch- 
ard-2 site, except in summer 1983 when survival of 
Deer Mice was significantly lower in the orchard 
than old field sites (Figure 3). 

Mean body mass of Deer Mice was the same or 
higher in orchard than old field sites during summer 
and winter periods (Figure 4). Only during winter 
1983-1984 did body mass of mice in orchard-2 appear 
lower than that in the old field sites. 


TABLE 2. Proportion of adult male and female Deer Mice in reproductive condition during breeding seasons each year for 
pooled data in replicate old field and orchard sites and results of Chi-square (y)analysis. Sample size (number of mice) in 
parentheses. Significant P in bold face. 


Old field Orchard Analysis 
Year ve PB. 
Males 
1982 0.62 (84) 0.79 (28) 2.60 0.11 
1983 0.67 (367) 0.95 (114) 34.87 <0.01 
1984 0.67 (253) 0.81 (86) 6.58 0.01 
1985 0.79 (150) 0.97 (39) qo <0.01 
Females 
1982 0.26 (69) 0.38 (42) eg 0.19 
1983 0.30 (209) 0.30 (106) 0.00 1.00 
1984 0.40 (182) 0.64 (96) 13.82 <0.01 
1985 0.47 (103) 0.61 (49) 2.84 0.09 


TABLE 3. Mean number of recruits (X) + 95% confidence intervals for Deer Mice during summer and winter periods in 
replicate old field and orchard sites. Sample size (n = number of trapping periods) in parentheses. 


Old field Orchard 

Period 1 2 1 2 
Summer 1982 X Syl Sel 4.6 5.6 

(7) 95% Cl 2.6 —7.7 6.2 — 20.1 1:27 8:0 2.2 -8.9 
Winter 1982-1983 X 4.0 6.6 2.8 2) 

(5) 95% CI 1.7-6.3 2.6 — 10.6 0.0 —5.6 1.2—3.2 
Summer 1983 X 6.7 10.0 49 5.6 

(9) 95% CI 2.0— 11.3 SO llsy(9) DAVIS) 2.2 —-8.9 
Winter 1983-1984 x BS) 3.5 1.9 3.8 

(8) 95% CI 1.5-9.5 169-4 0.8 — 2.9 1.9-5.6 
Summer 1984 X 9.9 SS) DS) 5.8 

(8) 95% CI 7.3 —12.4 2.5 —8.5 2.5—8.5 3.2 — 8.3 
Winter 1984-1985 X 3d 1S De), 0.7 

(6) 95% CI 0.6 — 6.7 -0.6 — 3.6 -1.0 —5.3 One? 
Summer 1985 X 7.8 3.4 3.9 2.0 

(9) 95% Cl 4.3—11.2 2.1-—4.8 1.9-—5.9 1.1-—2.9 
Winter 1985-1986 X 8.6 3.9 7.0 Dal 

(7) 95% CI 1.5 — 15.6 0.5 -— 7.3 eS P27/ 0.2 —4.1 


304 THE CANADIAN FIELD-NATURALIST Vol. 118 
F @ i O Orchard 1 
Peromyscus maniculatus orice” mn 
@ Old field 2 O Orchard 2 
| 
O 
2 
c LJ 
2 06 " 
© 
ne) 
1) 
of 
> 
= 04 if ? 
2 
S 
14) 
oO 
=> 
0.2 
0 
Summer Winter Summer Winter Summer Winter Summer Winter 
1982 1982-83 1983 1983-84 1984 1984-85 1985 1985-86 


FiGurE 3. Mean Jolly-Seber survival (per 28 days) + 95% confidence intervals for Deer Mice in replicate old field and 
orchard sites during summer and winter periods in the study. 


Yellow-pine Chipmunk populations 

Totals of 137 and 79 individual Yellow-pine Chip- 
munks were captured in the two old field and two 
orchard sites, respectively. Only three individual chip- 
munks were captured in the orchard-2 site during the 
study. Mean J-S trappability estimates ranged from 
30.8% to 63.8% in the old field sites, and from 37.1% 
to 58.3% in the orchard sites. 

Populations of Yellow-pine Chipmunks ranged in 
mean abundance/ha from 5.6 to 19.0 in old field sites 
and from 1.9 to 17.5 in the orchard-1 site (Table 4). 
Mean abundance of Yellow-pine Chipmunks was sig- 
nificantly (non-overlapping 95% confidence intervals) 
lower in the orchard-1 than old field sites in 1982 and 
1985 (Table 4). In terms of population changes over 
the study, Yellow-pine Chipmunk abundance/ha ranged 
from 1.0 — 27.0 in old field sites and from 1.0 — 46.3 
in the orchard-1 site (Figure 5). 

Mean number of Yellow-pine Chipmunk recruits 
was similar between the old field and orchard-1 sites 
(Table 4). Mean (+ SE) Jolly-Seber survival rates 
averaged 0.78 + 0.05 (summer) and 0.96 + 0.02 (win- 


ter) in the old field-1 site and 0.71 + 0.05 (summer) 
and 0.91 + 0.06 (winter) in the orchard-1 site. 


Discussion 
Deer Mouse and Yellow-pine Chipmunk populations 
The pattern of abundance of Deer Mice in our old 
field sites was somewhat different from that recorded 
in other old field and perennial grassland studies of 
Peromyscus and Microtus. Grant (1972) and Baker 
(1968) suggested that the Deer Mouse suffers from 
competition from microtine rodents in perennial grass- 
lands, and is forced to live in woodlands. However, 
Deer Mouse densities in our studies reached annual 
peaks ranging from 30.8 to 69.3 animals/ha in the pres- 
ence of relatively high numbers of Montane Voles 
(Sullivan et al. 2003). Experimental studies by Grant 
(1971) and Redfield et al. (1977) concluded that Micro- 
tus Outcompete Peromyscus in grassland habitats; how- 
ever, explanations for this process were not given. 
Conversely, other studies concluded that competitive 
interactions among microtine rodents and Deer Mice 
were unimportant (Gilbert and Krebs 1984; Galindo 


2004 SULLIVAN, SULLIVAN, AND HOGUE: POPULATION DYNAMICS OF DEER MICE AND CHIPMUNKS 305 


® Old field 1 O Orchard 1 
Old field 2 O Orchard 2 


Peromyscus maniculatus 


30 
35 
43 
S 86 
21 
my CD oA 57 
a 97 141} J 169 ,, 13 68 478 
= t 10440) ze 
44 36 
> mA 96 90 
3 72 158 
2 + ' 220 
87 

= if 355 ; 
= { 
= 2) 

15 


Summer Winter Summer Winter Summer Winter Summer Winter 
1982 1982-83 1983 1983-84 1984 1984-85 1985 1985-86 


FiGuRE 4. Mean body mass (g) + 95% confidence intervals for Deer Mice in replicate old field and orchard sites during 
summer and winter periods in the study. 


TABLE 4. Mean abundance (X) and mean number of recruits per ha (X) (+ 95% confidence intervals) for Yellow-pine 
Chipmunks during non-hibernation periods in replicate old field and orchard sites. Sample size (x = number of trapping 
periods) in parentheses. 


Attribute and Old field Orchard 

period 1 2 1 2 
Abundance 

1982 X 5.6 7.9 1.9 - 
(7) 95% CI 3.6 — 7.7 3.8 — 12:0 0.7 — 3.0 - 
1983 X 8.1 11.8 9.3 0.1 
(13) 95% Cl 5.3 — 10.9 9.8 — 13.8 2.2 — 16.4 = 
1984 X 19.0 8.9 TES) 0.2 
(11) 95% CI 15.2 — 22.9 6.2 — 11.6 8.7 — 26.2 - 
1985 X 14.5 9.8 3.6 - 
(10) 95% Cl 12.1 — 16.9 6.2 — 13.4 1.9-—5.4 — 
Recruits 

1982 X 1.1 Da 0.6 - 
(7) 95% CI 0.5 — 1.8 0.1-—5.3 -0.2 — 1.3 — 
1983 X eS) 12 2.3 0.1 
(13) 95% Cl 0.1 — 3.0 0.0 — 2.4 0.3 — 4.3 - 
1984 x De) 0.7 3.4 0.2 
(11) 95% CI E2339 -0.1-1.5 0.6 —6.1 ~ 
1985 X 1S 2.1 0.4 - 


(10) 95% CI 0.7 — 2.3 0.5 — 3.7 -0.3-1.1 = 


306 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Tamias amoenus 


50 


N () as 
(o) (~) oO 


Number of animals/ha 


=x 
oO 


--Old field 1 
-# Old field 2 
< Orchard 1 


J S NJ MM:J SON JOM.M J°S.N JOM Mod -S NLS MiGM 


1982 1983 


1984 1985 1986 


FiGuRE 5. Population densities (Jolly-Seber) per ha of Yellow-pine Chipmunks in two old field sites and one orchard site 
during the study. Shaded bars indicate winter periods. Months of year are represented by J = July; S = September; N 


= November; J = January; M = March; M = May. 


and Krebs 1985). Sullivan and Krebs (1981) reported 
persistence of Deer Mice with Microtus spp. in a grass- 
land habitat. Our densities of Deer Mice were as high 
as reported in other studies of perennial grassland hab- 
itats and orchards (Sullivan and Krebs 1981; Sullivan 
et al. 1998). 

This study is the first detailed investigation of the 
population dynamics of non-target rodent species in 
orchards treated with rodenticides for vole control. 
Populations of Deer Mice in orchard sites showed con- 
sistent annual changes in abundance averaging 15.0 
animals/ha in summer and 10.1 animals/ha in winter 
periods. These values were similar to those reported 
for Montane Voles in these same orchards at 16.5 ani- 
mals/ha in summer and 21.1 animals/ha in winter (Sul- 
livan et al. 2003). Clearly, Deer Mice seemed produc- 
tive in the orchard environment, having longer breed- 
ing seasons than in the old field sites, and comparable 
survival of young animals. Winter breeding in Deer 
Mice has been reported, particularly after substantial 
mast crops in forests (Wolff 1996) or in supplemental 
food studies (Taitt 1981). Food resources appeared 


sufficient to support breeding mice during winter peri- 
ods and at body masses comparable to old field popu- 
lations during most periods. These orchards received 
regular inputs of fertilizer and irrigation, and hence 
likely produced relatively “rich” habitats in terms of 
plant production and invertebrate biomass. Mean 
overall survival of Deer Mice in orchards was poor, in 
at least one of the sites, relative to old field populations. 
A lack of vegetative cover may have contributed to 
greater predation by raptors, weasels (Mustela spp.), 
and Coyotes (Canis latrans). 

Our orchards were situated in a mosaic of different 
varieties and age classes of orchards and vineyards 
with adjacent natural habitats of sagebrush and Pon- 
derosa Pine (Pinus ponderosa). As discussed by Sul- 
livan et al. (1998) for their study in this same mosaic, 
it was possible that our relatively high numbers of 
Deer Mice (and Yellow-pine Chipmunks) in orchards 
may have represented transients or animals who lived 
in nearby sage-pine areas. However, a comprehensive 
analysis of transient and resident animals suggested 
strongly that these rodents very likely lived in the actual 


2004 


orchard units (Sullivan et al. 1998). There was no rea- 
son to assume that our animal populations behaved 
differently, and hence our population estimates were 
considered accurate. 

The prediction of the hypothesis that population 
dynamics of Deer Mice and Yellow-pine Chipmunks 
would be enhanced in old field compared with orch- 
ard habitats is partially accepted. Abundance, recruit- 
ment, and overall survival of Deer Mice were higher 
in old field than orchard sites. However, proportion 
of breeding animals and length of breeding seasons 
were greater in the orchard sites. Early juvenile sur- 
vival and body mass were similar between sites for 
Deer Mice. Similarly, the general lack of differences 
in abundance (except in 1982 and 1985) and recruits 
for Yellow-pine Chipmunks between sites (albeit for 
one replicate only) also contradicted our prediction. 

These rodent species should be able to maintain 
their population levels in association with Microtus 
in these habitats. Traditional methods of vole control 
(rodenticides) seem to have had little effect on these 
two non-target species in the orchard sites. The higher 
abundance of Deer Mice, and sometimes Yellow-pine 
Chipmunks, in the old field than orchard sites was 
more likely a function of habitat quality than expo- 
sure to rodenticides. Consequently, Deer Mice and 
Yellow-pine Chipmunks may assist integrated pest 
management in orchards because both species per- 
sisted in this managed habitat and their consumption 
of seeds and invertebrates could provide some degree 
of assistance in reducing weed and insect pests. In 
addition, as prey species, they may attract a greater 
number of predators to these sites. A total of six small 
mammal species was recorded in these orchards, 
which suggests that these agro-ecosystems and adja- 
cent natural lands help contribute to a diversity of 
habitats in this agrarian landscape. 


Acknowledgments 

We thank the Research Branch of Agriculture Can- 
ada and the Applied Mammal Research Institute for 
financial support; the study area and facilities provided 
by the Summerland Research Station; and field assis- 
tants W. Kaiser, J. Gareau, B. Jones, J. Krebs, and C. 
Robinson. 


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Received 11 August 2002 
Accepted 24 December 2004 


Density and Survival of Lady Beetles (Coccinellidae) in Overwintering 
Sites in Manitoba* 


W. J. TURNOCK and I. L. WISE 


Cereal Research Centre, Agriculture and Agri-Food Canada, 195 Dafoe Road, Winnipeg, Manitoba, Canada R3T 2M9 
Turnock, W. J., and I. L. Wise. 2004. Density and survival of Lady Beetles (Coccinellidae) in overwintering sites in Manitoba. 
Canadian Field-Naturalist 118(3): 309-317. 


The densities of lady beetles, Coccinellidae, overwintering as adults (adults per m7) in leaf litter collected in late October for 
two years in a beach-ridge forest on the south shore of Lake Manitoba were 56.4 for the Thirteen-spotted Lady Beetle, 
Hippodamia tredecimpunctata (Say), 38.3 for the Seven-spotted Lady Beetle, Coccinella septempunctata (L.), 7.7 for the 
Transverse Lady Beetle, Coccinella transversoguttata richardsonii Brown, 1.6 for the Convergent Lady Beetle, Hippodamia 
convergens Guerin, and 0.6 for the Parenthesis Lady Beetle, Hippodamia parenthesis (Say). The mean overwintering survival 
for these species was 0.254, 0.036, 0.023, 0.0, and 0.0, respectively. The density of overwintering coccinellids was highest 
near the margins of the forest, particularly on the beach side, where beetles from shore appear to have entered the forest. The 
mean density over 3 years (2.9 per m7) of all coccinellid species in November in the litter under a remnant grove of riverbank 
forest in Winnipeg, was lower than in the beach-ridge forest (104.8 per m?), but their survival (0.460) was higher than in the 
beach-ridge forest (0.154). More species of coccinellids were found in the samples from the riverbank forest than from the 


beach-ridge forest. 


Key Words: Lady Beetles, Coccinellidae, leaf litter, beach-ridge forest, overwintering sites, Lake Manitoba, Red River. 


The overwintering sites of north-temperate zone coc- 
cinellids include leaf litter, grass tufts, bark crevices, 
stone piles, and rock clefts (Hodek and Honék 1996). 
Adults of some species migrate long distances to hiber- 
nate at high elevations, and even those which move 
only locally seem to be attracted to even slightly higher 
elevations and vertical silhouettes. In Bohemia and 
Moravia, Coccinella septempunctata L. (C7) prefers 
elevated places, but also hibernates in refuges among 
cultivated areas and does not migrate long distances 
(Hodek 1960). In southern Manitoba, where much of 
the land is flat and cultivated, it is generally believed 
that lady beetles overwinter in the leaf litter under trees 
in field shelterbelts, farmsteads and urban plantings, 
along the shores of streams and lakes, and in “bluffs” 
(groves of trees on uncultivated land). The species oc- 
currence, density, and survival of coccinellids in most 
of these overwintering sites have not been studied. 

Large numbers of pre-hibernating coccinellids are 
washed ashore in some years on Manitoba lakes, usu- 
ally in late August to early October (Lee 1980; Turnock 
et al. 2003). These beetles usually fly away soon after, 
to complete their search for hibernating sites, and do 
not often walk into the adjacent beach-ridge forest. 
Once, on 1 November 1978, beetles (mainly H. con- 
vergens) were washed ashore, but subsequent cold 
weather forced them to remain near the beach, where 
few survived the winter (Turnock and Turnock 1979). 

In this paper, the density, distribution, species com- 
position, and overwintering survival of coccinellids in 
the litter under a beach-ridge forest are compared with 


the density and survival of coccinellids in the litter 
under a riverbank forest. 


Methods 

Beach-Ridge Forest. The abundance and species 
composition of coccinellids overwintering in the sur- 
face litter and soil under a beach-ridge forest on the 
south shore of Lake Manitoba at the Delta Marsh Field 
Station (University of Manitoba), 50°11'N, 98°23'W, 
about 120 km WNW of Winnipeg, were recorded. The 
shore is a sand beach with a forested barrier-beach 
ridge formed by the reworking of alluvial sand depos- 
its. This beach ridge, about 60 m wide, supports a 
mature deciduous forest of Manitoba Maple (Acer 
negundo L.), Green Ash (Fraxinus pennsylvanica 
Marsh.), Plains Cottonwood (Populus deltoides Marsh.), 
and Peachleaf Willow (Salix amagdaloides Anderss.) 
(Kenkel 1986). The ridge has its highest elevation 5- 
10 m from the top of the foreshore and slopes down 
about 50 m to the marsh. A road about 10 m wide along 
the south edge of the beach ridge separates the forest 
from the marsh. Samples were taken along five tran- 
sects, 20 m apart, perpendicular to the beach. Six sam- 
ples were taken along each transect at 1, 6, 11, 21, 31, 
and 41 m from the top of the foreshore, where the 
beach ended and the beach-ridge forest began. A 
sample of the leaf litter and soil above the sandy par- 
ent material, 0.25 m? and 5-10 cm deep, was collect- 
ed at each of the 30 locations. Samples were collected 
after beetle flight had ceased, on 22 October 1992 and 
27 October 1993. Each sample was bagged, sorted by 


*Contribution Number 1832 from the Cereal Research Centre, Agriculture and Agri-Food Canada, 195 Dafoe Road, Winnipeg, Manitoba, 


Canada R3T 2M9. 


309 


310 


hand to remove the beetles, and the species and num- 
ber of coccinellids were recorded. 

The numbers and species of coccinellids emerging 
in the spring were determined from cone traps, each 
covering a surface area of 0.1028 m* (Turnock et al. 
1987). One trap was set into the litter at each of the 
30 locations, near the place where a litter sample had 
been collected the previous autumn. The traps were 
set out at each location in late April or May, depend- 
ing on the disappearance of snow, in 1993, 1994, and 
1995. The traps at the apex of the cones were emptied 
at regular intervals, and the coccinellids identified to 
species and counted. 

Riverbank Forest. The overwintering of coccinellids 
in a remnant grove of riverbank forest along the Red 
River at the University of Manitoba, Winnipeg, Mani- 
toba, was determined from records of the insects found 
in litter samples and emergence traps used to study 
the overwintering of flea beetles (Chrysomelidae) (Tur- 
nock et al. 1987). The grove of trees, about 1.3 ha in 
area, included Bur Oak (Quercus macrocarpa Michx.), 
American Elm (Ul/mus americana), Manitoba Maple, 
Green Ash, Black Ash (Fraxinus nigra Marsh.), and 
Basswood (Tilia americana L.). In this grove, 50 litter 
samples, each 0.25 m7, <10 cm deep, were collected in 
the autumn of the years 1980-1982, and in the springs 
of 1980, 1981, and 1983. The samples were taken at 
10 m intervals along five transects running north to 
south through the grove. Spring emergence was sam- 
pled by 100 cone traps, each covering 0.1028 m/, in 
1980 and 1981. Two traps were located at 10 m inter- 
vals along each transect, one on each side of the site 
of a litter sample, as described by Turnock et al. (1987). 
For coccinellids, only the total number collected in 
each sampling period was recorded, except in the litter 
samples of 3 November 1982, where the species were 
identified but their numbers were not recorded. 


Results 

Beach-Ridge Forest. The dominant species in all 
collections were H. tredecimpunctata, C. septempunc- 
tata, and Coccinella transversoguttata. In the autumn 
litter samples in 1992 (n = 653) and 1993 (n = 918) 
coccinellid relative abundances were: 0.54/0.54 (H. 
tredecimpunctata); 0.36/0.37 (C. septempunctata); 0.08/ 
0.07 (C. transversoguttata); 0.01/0.02 H. convergens; 
and 0.004/0.001 H. parenthesis (Say). In the spring 
emergence samples, the relative abundances of each 
species in 1993/94/95 were: H. tredecimpunctata — 
0.95/0.78/0.92; C. septempunctata — 0.04/0.19/0.06; 
and C. transversoguttata — 0.02/0/0. Neither H. conver- 
gens nor H. parenthesis were found in the spring sam- 
ples. The distribution of numbers per sample of H. 
tredecimpunctata and C. septempunctata was skewed 
in both the autumn litter samples and the spring emer- 
gence samples, with most samples containing 0, 1, or 
2 beetles (Figure 1). 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


The mean density per m* of coccinellids in the 
autumn was 87.1 in 1992 and 122.4 in 1993, and 8.5 
and 11.7 in the following springs. The density of coc- 
cinellids in the autumn was generally higher on the 
margins of the forest, but it varied among species 
(Figure 2). The numbers of H. tredecimpunctata were 
highest near the beach margin in 1993, but they were 
higher near the marsh margin in 1992. The density of 
both C. septempunctata and C. transversoguttata was 
higher near the beach margin in both years, with only a 
slight increase along the marsh margin. These patterns 
suggest that most of the C. septempunctata and C. 
transversoguttata found in the litter near the beach were 
from aggregations that had washed ashore (Turnock et 
al. 2003). Hippodamia tredecimpunctata seems mainly 
to have come from the beach in 1993, but in 1992 
more appeared to have come across the marsh to the 
edge of the forest. 

The proportion of living beetles in the autumn litter 
samples differed little between years, 81.3% in 1992 
(n = 653) and 81.4 % (n= 913) in 1993 (Table 1). The 
mean proportion of living beetles, by species for both 
years, was: H. tredecimpunctata = 0.963 (n = 847); 
C. septempunctata = 0.729 (n = 575); C. transversogut- 
tata = 0.250 (n = 116); H. convergens = 0.458 (n = 24); 
and H. parenthesis = 0.500 (n = 4). Survival of all spe- 
cies was lowest adjacent to the beach and increased 
toward the edge of the marsh (Figure 3). The decreased 
survival in the forest edge near the beach was prob- 
ably caused by the burial of overwintering beetles by 
sand blown off the beach by strong winds. 

Coccinellids emerged from the leaf litter during May 
and early June in traps throughout the beach-ridge. 
Most of the beetles (98%, n = 187) were of two spe- 
cies, H. tredecimpunctata and C. septempunctata. The 
mean density per m? of emerging coccinellids in 1993 
to 1995 was: 17.5, 9.1, 27.9 for H. tredecimpunctata; 
0.65, 2.3, 1.9 for C. septempunctata; and 0.32, 0, 0 for 
C. transversoguttata. The pattern of higher densities 
near the margins of the forest was less prominent in 
the spring than in the autumn (Figures 2 and 4). The 
density of emerging beetles was <20 per m? except at 
distances from the beach of | m in 1995, 31 m in 1994, 
and 41 m in 1993. 

The mean overwintering survival (density spring/ 
density autumn) in the beach-ridge forest for H. tre- 
decimpunctata was 0.370 in the winter of 1992-1993, 
and 0.139 for 1993-1994 (Table 1). The survival ratios 
over the same two winters for C. septempunctata 
were (0.021 and 0.051, and for all coccinellids, 0.212 
and 0.096. The survival of H. tredecimpunctata was 
quite variable with position within the forest (Figure 5). 
Overwintering survival of all species was very low at 
1 m from the beach margin of the forest, where drifting 
sand often covered the leaf litter. However, the over- 
wintering survival did not significantly differ among 
years, density of coccinellids in the autumn, or distance 


2004 TURNOCK and WISE: DENSITY AND SURVIVAL OF LADY BEETLES 31] 


Coccinella septempunctata 


Mmm 1992 
Gas%] 1993 


Frequency 


60 80 100 


Hippodamia tredecimpunctata 


Frequency 


100 


Coccinella transversoguttata 


Frequency 


80 100 


0 20 40 60 


Beetles per sample 


FIGURE 1. Frequency distribution of numbers of coccinellids, Hippodamia tredecimpunctata, Coccinella septempunctata, 
and Coccinella transversoguttata, per sample of litter (0.25 m? x 5-10 cm deep) from the floor of the beach-ridge 
forest at the Delta Marsh Field Station, Manitoba, in late October of 1992 and 1993. n = 60 for each species. 


SH 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE 1. The mean numbers (N) per m? in autumn litter samples and spring emergence samples, the proportion of living 
beetles in the autumn samples, and the overwintering survival (density spring emergents/density in autumn), for Coccinella 


septempunctata (C7), Hippodamia tredecimpunctata (H13), 


C. transversoguttata (CT), H. convergens (HC), and H. 


parenthesis (HP), and all coccinellids in the beach-ridge forest at the Delta Marsh Field Station. 


Species 1992 1993 
Autumn Spring 
H13 N/m? 47.3 IES 
Survival 0.95 0.37 
N 355 54 
(Oy N/m? Si1k3 0.65 
Survival 0.72 0.02 
N 235 2 
CT N/m? 6.9 0.32 
Survival 0.37 0.05 
N 52 1 
HC N/m2 101 0 
Survival 0.38 0 
N 8 0 
HP N/m? 0.4 0 
Survival 0.67 0 
N 3 0 
All N/m? 87.1 18.5 
Survival 0.81 0.21 
N 653 57 


1993 1994 1995 
Autumn Spring Spring 
65.6 9.1 27.9 
0.97 0.14 ~ 
492 28 86 

45.3 ep) 129 
0.73 0.05 - 
340 q 6 
8.5 0 0 
0.16 0 - 
64 0 0 
25k 0 0 
0.5 0 - 
16 0 0 
0.13 0 0 
1 0 - 

0 0 
122.4 11.7 30.5 
0.81 0.1 — 

913 36* 94% 


*Includes one specimen of Anistosticta bitriangularis in 1994 and two specimens of Calvia quatuordecimguttata in 1995. 


from the beach (analysis of variance, proc glm, SAS 
Inst. Inc. <www.sas.com>). The high survival of H. tre- 
decimpunctata at several locations likely reflects the 
placing of the emergence traps over local aggrega- 
tions of beetles in the litter, as does a single record of 
high survival of C. septempunctata. 


Riverbank Forest. The litter samples collected on 3 
November 1982 contained nine species of coccinel- 
lids: Stethorus punctum (LeConte), Scymnus (Pullus) 
brullei Muls., Hyperaspis benedetti (Say), Chilocorus 
stigma Say, H. tredecimpunctata, Anatis labiculata 
(Say), Adalia bipunctata L., C. transversoguttata, and 
Psyllobora virginimaculata (Say). The mean density 
of all coccinellid species in the autumn litter collec- 
tions of 1980-1982 (n = 110) was 2.9 per m? (2.0- 
4.3), vs. 4.4 (1.5- 10.4) in the spring litter collections 
(n = 172) of 1980, 1981, and 1983. Overwintering 
survival, calculated by dividing the number of living 
beetles in the spring litter samples by the total num- 
ber of beetles in the autumn samples, was 76% for the 
winter of 1980-1981 and 43% for the winter of 1982- 
1983. Survival during the winter of 1980-1981 was 
lower, 49%, when the spring density, 0.97 per m*, was 
based on captures in the emergence traps (n = 10). 


Discussion and Conclusions 

The studies of coccinellid density and survival in 
three overwintering sites in southern Manitoba differ 
in location, years, and sampling design. However, the 
results of some comparisons may be helpful to future 
investigators of site selection and survival by overwin- 


tering coccinellids. In the autumn, more overwintering 
lady beetles were found in a beach-ridge forest on 
the shores of Lake Manitoba in 1992 and 1993 than in 
a riverbank forest in Winnipeg in 1980 to 1981. The 
difference in the density between the two sites may 
have been related to differences in the populations of 
lady beetles in the years sampled. Another factor could 
be the availability of overwintering sites in the two areas. 
The beach-ridge forest was located in an intensively- 
farmed area with few treed areas, whereas the river- 
bank forest was near suburban Winnipeg, where trees 
were more abundant. Beetles washed ashore on the 
beach contributed to the number overwintering in the 
beach-ridge forest in 1993, but in 1992 more beetles 
seem to have entered the forest from the side away 
from the beach. 

The beach-ridge forest, despite higher initial den- 
sities, had fewer species and fewer beetles emerging 
in the spring than in the riverbank forest, because of 
lower overwintering survival. Overwintering survival 
by beetles may be related to their need for moisture 
(Hodek and Honék 1996). The litter under the river- 
bank forest was thicker and lay upon a clay soil, thus 
providing a moister environment than the thinner litter 
on a sandy soil under the beach-ridge forest. 

The limited sampling by Turnock and Turnock 
(1979) on the shores of Lake Manitoba, following a 
very late flight (1 November) and aggregation of 
beetles, indicated few beetles survived the winter (2% 
of 1002 per m? trapped in the beach debris). Even this 
level of survival is surprising, because beach debris 
usually is pounded and buried by high water, large 


2004 TURNOCK and WISE: DENSITY AND SURVIVAL OF LADY BEETLES 


Beetles per m? 


Beetles per m? 


Beetles per m? 


30 40 50 


0 10 20 


Distance from forest edge (m) 


FiGurE 2. Mean density per m? of Hippodamia tredecimpunctata, Coccinella septempunctata, Coccinella transversoguttata, 
and all coccinellids, in forest litter samples collected at various distances from the forest edge next to the beach in 


late October 1992 and 1993 at the Delta Marsh Field Station. 


314 THE CANADIAN FIELD-NATURALIST Vol. 118 


Proportion alive 


Proportion alive 


All species 


Proportion alive 


0 10 20 30 40 50 


Distance from forest edge (m) 


FicuRE 3. Proportion of live individuals of Hippodamia tredecimpunctata (H13), Coccinella septempunctata ( (C7). 
Coccinella transversoguttata (CT), and all coccinellids, in litter samples taken at different distances from forest edge 
next to the beach in late October of 1992 and 1993. 


TURNOCK and WISE: DENSITY AND SURVIVAL OF LADY BEETLES 


2004 


—e— H13 
-O::- C7 

N —-w— All species 
= —. 
= 

(oy 

[ou 

0) 
a) 

x) 

® 

faa) 

N 

E 

= 

(or) 

Q. 

o 

v 

zr) 

® 

a 

N 

i= 

= 

® 

a 

72) 

& 

© 

® 

a 


Distance from forest edge (m) 


FiGure 4. Mean density per m? of Hippodamia tredecimpunctata (H13), Coccinella septempunctata (C7), and all coccinel- 
lids, in emergence trap samples collected at different distances from the forest edge next to the beach. The samples 


were collected from emergence cones placed over the litter in May of 1993, 1994, and 1995. 


316 


Survival 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


H13-1992/3 
H13-1993/4 
C7-1992/3 
C7-1993/4 


Distance from forest edge (m) 


FiGurE 5. Mean overwintering survival ratios for Hippodamia tredecimpunctata (H13), and Coccinella septempunctata (C7), 
for the winters of 1992-1993 and 1993-1994 in forest litter on the beach ridge at different distances from the forest 


edge next to the beach at the Delta Marsh Field Station. 


waves, and ice chunks from violent autumn and spring 
storms. In this study the top of the foreshore, where a 
covering of aspen leaves had accumulated on the sand, 
had 74 beetles per m* and overwintering survival was 
53%. This location is not likely to be a suitable over- 
wintering site in most years because of high water 
and waves. The grassy backshore, with little litter, had 
28 beetles per m’, of which 18% survived the winter. 
The foreshore of the sandy beaches and even the shal- 
low litter in the sparse beach-ridge forest of Trembling 
Aspen, Populus tremuloides Michx., do not appear to 
be selected by coccinellids for overwintering. Excep- 
tions can occur when beetles washed ashore after un- 
usually late autumn flights aggregate on the beach when 
subsequent temperatures are too cool to allow flight, 
but warm enough to allow limited walking up the beach 
to the backshore. In these cases the beach-ridge forest, 
even if unsuitable, is mainly a “last-chance” overwin- 
tering site for these beetles. 

Four species of coccinellids, H. tredecimpunctata, 
C. septempunctata, C. transversoguttata, and H. con- 
vergens (in order of relative abundance) were found 
in both years in autumn litter samples in the beach- 


ridge forest, but only the first two species were found 
in the spring emergence samples in all three years. 
More H. tredecimpunctata (14 and 38%) survived over 
winter than C. septempunctata (3 and 5%) or C. trans- 
versoguttata (5 and 0%). This level of survival of C. 
septempunctata is much lower than the 97% survival 
of C. septempunctata that had been covered with litter 
in experimental cages at Edmonton, Alberta (Ryan and 
Acorn 1999). 

Although in Europe C. septempunctata does not 
migrate long distances to hibernate (Hodek 1960), large 
flights of this species occur in both autumn and spring, 
as shown by the numbers of this species that are found 
in the aggregations washed ashore on the shores of 
the lakes (Turnock et al. 2003). 

Southern Manitoba is mostly cultivated and has a 
flat terrain, so potential overwintering sites are limited 
to leaf litter under trees. These habitats probably are 
both attractive and suitable for the hibernation of coc- 
cinellids, but without information on the preferred over- 
wintering sites in this area, the true level of overwinter- 
ing survival by native and introduced species cannot 
be compared. 


2004 


Acknowledgments 

The statistical advice by S. Woods, technical assis- 
tance of R. J. Bilodeau, and the help given by the staff 
of the Delta Marsh Field Station (University of Mani- 
toba) in providing facilities is gratefully acknowledged. 
This is Publication Number 305 from the Delta Marsh 
Field Station (University of Manitoba). 


Literature Cited 

Hodek, I. 1960. Hibernation-bionomics in Coccinellidae. 
Acta Societis Entomolicae Cechosloveniae 57(1): 14-17. 
[In Czech with English “Discussion and Conclusions” }. 

Hodek, I., and A. Honék. 1996. Ecology of Coccinellidae. 
Series Entomological. Volume 54. Kluwer Academic Pub- 
lishers. Dordrecht. 464 pages. 

Kenkel, N. C. 1986. Vegetation structure and dynamics of the 
barrier-beach ridge at Delta, Lake Manitoba. Annual Report 
of the University of Manitoba Delta Marsh Field Station 
21: 61-83. 


TURNOCK and WISE: DENSITY AND SURVIVAL OF LADY BEETLES 317 


Lee, R. E. 1980. Aggregations of lady beetles on the shores 
of lakes. American Midland Naturalist 104; 295-304. 

Ryan, S. E., and J. H. Acorn. 1999. Overwintering survival 
of the seven-spot ladybug (Coccinella septempunctata) in 
Edmonton. Blue Jay 57: 97-100. 

Turnock, W. J., and R. W. Turnock. 1979. Aggregations of 
lady beetles (Coleoptera: Coccinellidae) on the shores of 
Lake Manitoba. Manitoba Entomologist 13: 21-22. 

Turnock, W. J., R. J. Lamb, and R. J. Bilodeau. 1987. Abun- 
dance, winter survival, and spring emergence of flea beetles 
(Coleoptera: Chrysomelidae) in a Manitoba grove. The 
Canadian Entomologist 119: 419-426. 

Turnock, W. J., I. L. Wise, and F. O. Matheson. 2003. Abun- 
dance of native coccinellines (Coleoptera: Coccinellidae) 
before and after the appearance of Coccinella septem- 
punctata. The Canadian Entomologist 135: 391-404. 


Received 9 August 2002 
Accepted 24 December 2004 


Description of Age-O0 Round Goby, Neogobius melanostomus Pallas 
(Gobiidae), and Ecotone Utilisation in St. Clair Lowland Waters, 
Ontario 


JOHN K. LESLIE and CHARLES A. TIMMINS 


Great Lakes Laboratory for Fisheries and Aquatic Sciences, Department of Fisheries and Oceans, 867 Lakeshore Road, Burling- 
ton, Ontario L7R 4A6 Canada 


Leslie, John K., and Charles A. Timmins. 2004. Description of age-O Round Goby, Neogobius melanostomus Pallas (Gobiidae), 
and ecotone utilisation in St. Clair lowland waters, Ontario. Canadian Field-Naturalist 118(3): 318-325. 


Early developmental stages and ecotone utilisation of the non-indigenous Round Goby, Neogobius melanostomus (Pallas, 1811), 
are described and illustrated. Fish (5-40 mm) were collected in coarse gravel, rocks and debris in the St. Clair River/Lake 
system, Ontario, in 1994-2000. The Round Goby hatches at about 5 mm with black eyes, flexed urostyle, and developed fins 
and digestive system. Distinguishing characters include large head, dorsolateral eyes, large fan-shaped pectoral fins, two dorsal 
fins, fused thoracic pelvic fins and a distinct black spot on the posterior of the spinous dorsal fin. Modal counts for preanal, 
postanal, and total myomeres were 12, 19, and 31, respectively. 


Key Words: Round Goby, Neogobius melanostomus, St. Clair aquatic ecosystem, age-0, morphometry, habitat, Ontario. 


Gobiidae, the largest family of marine fishes, has 
been found in fresh waters of all continents (Nelson 
1984) except Antarctica. None of 68 species endemic 
to North America is native to the Great Lakes, where 
two introduced species, the Round Goby, Neogobius 
melanostomus (Pallas, 1811) and the Tubenose Goby, 
Proterorhinus marmoratus (Pallas, 1811), recently be- 
came established. These fishes originate in the Ponto- 
Caspian region (Caspian, Azov and Black seas and Sea 
of Marmara). International ships’ ballast was assumed 
the most probable means by which Round Goby eggs 
and/or postembryonic stages were transferred from 
the Black Sea to the Great Lakes (Crossman et al. 
1992; Jude et al. 1992). Since its arrival in the mid- 
late 1980s the relatively innocuous Tubenose Goby 
has not proliferated beyond the St. Clair-western 
Lake Erie ecosystem (Leslie et al. 2002). In contrast, 
the aggressive, eurytopic Round Goby has diffused 
extensively and is now one of the most abundant fish 
in the Great Lakes basin. Apparently, repetitive spawn- 
ing and habitat requirements of this fish conflict with 
our native species (Dubs and Corkum 1996), some of 
which they may supplant. 

The Round Goby is a guarding cavity spawner with 
a life span of 4-5 years (Miller 1986). Spawning sea- 
son is typically from spring to late summer. Rocks, 
gravel, solid objects with crevices and submersed plants 
are used for egg deposition (Miller 1986; Skora 1997*). 
Whereas literature abounds on the ecology of adults, 
information on postembryonic fish is rare. Eggs and 
early developmental stages in southern Russia and 
Ukraine have been described by Moskal’kova (1967), 
Kalinina (1976), and Koblickaja (1981). This report 
presents information on aspects of taxonomy and eco- 
tone utilisation of age-O0 Round Gobies in the Great 


Lakes and considers possible effects the species may 
have on the aquatic community in general and native 
fishes in particular. 


Study Area 

Eggs and age-0 fish were collected at numerous 
locations at the shore of the St. Clair River, Ontario, 
various sites along the southeastern shore of Lake St. 
Clair, and in Duck Creek (42°17'N, 82°35'W), a low 
gradient stream tributary to Lake St. Clair (Figure 1). 
Duck Creek is a disturbed system draining a small 
area in one of the most highly cultivated agricultural 
regions in Canada. It is approximately 9 km long and 
10-15 m wide near its mouth, where most specimens 
were collected. Mid-stream depth ranges from 1.2 to 
1.6 m. In April to November 1995, mean conductivity 
was 487 uS/cm and Secchi disc depth 0.3 + 0.1 m, 
reflecting high suspended particulate load due to run- 
off from cropland. Substrate at the sampling site is 
mainly alluvium, with clay and scattered debris of 
human origin at the base of 1-4 m high stream banks. 
However, age-O Round Gobies were found only on a 
small gravelled area at the base of a railway bridge. 

Duck Creek has a large diverse, albeit transient, fish 
assemblage of at least 41 species (Leslie and Timmins 
1998) dominated by Gizzard Shad Dorosoma cepedi- 
anum, Bluntnose Minnow Pimephales notatus, Spottail 
Shiner Notropis hudsonius, and Bluegill Lepomis 
macrochirus. Submersed plants near the collection site 
include Pondweed Potamogeton spp., Eelgrass Vallis- 
neria americana, Canada Waterweed Elodea cana- 
densis, Eurasian Milfoil Myriophyllum spicatum, and 
Curly Pondweed Potamogeton crispus. Arrowhead 
Sagittaria sp., Sedge Carex sp., Cattail Typha spp., and 
Bulrush Scirpus spp. were abundant emergent species. 


318 


2004 


Michigan Lake Huron 
ichig 


20 kn 


Lake St. Clair 


© Duck Creek 


FiGure 1. Main sites where age-O Neogobius melanostomus 
were collected in the St. Clair River-Lake ecosystem 
in 1994-2000: LGS (Lambton Generating Station) and 
Duck Creek. 


Round Goby eggs and age-0 developmental stages 
as well as numerous co-occurring fishes were found 
at various locations at the shore of the St. Clair River, 
mainly in dilute condenser cooling water discharge 
of Lambton Generating Station (LGS) (42°47'N, 
82°22'W). Random fish collections took place at LGS 
(Figure 1) each year between April and November 
1994-2000 over a shoreline stabilised with debris, 
limestone boulders and rocks 0.3-1.0 m in longest 
dimension. This formation extends 4 m from shore on 
a natural substrate of sand. Most common fish species 
collected with age-O Round Gobies in the St. Clair 
River include age-O0 White Sucker Catostomus com- 
mersoni, Alewife Alosa pseudoharengus, Brook 
Silverside Labidesthes sicculus, Rainbow Darter Ethe- 
ostoma caeruleum, lowa Darter Etheostoma exile, 
Johnny Darter Etheostoma nigrum, Spottail Shiner, 
and Emerald Shiner Notropis atherinoides. Of these 
species, guarding speleophils Rainbow Darter, Iowa 
Darter, and Johnny Darter utilise common ecotones 
and probably compete with gobies for food and shelter. 


Methods 

A bulging larval fish beach seine (4 m long, 1 m 
wide, mesh 0.4 mm) was necessary for capture of 
recently hatched gobies, which only emerged from pro- 
tective crevices when the bottom of the seine dis- 


LESLIE and TIMMINS: AGE-O ROUND GoBy IN ONTARIO 


319 


turbed the substrate. Two flat seines, 3 m long, | m 
wide (3 mm mesh), and 6 m long, | m wide (6 mm 
mesh) were used to sample young and adult gobies 
and co-occurring species. All seine hauls took place 
repeatedly in contact with gravel at the base of a rail- 
way bridge over Duck Creek. 

In the St. Clair River, clusters of eggs attached to 
rocks or human artefacts such as discarded sewer 
pipe were preserved intact with 5-10% formalin and 
enumerated in the laboratory. Water temperature, spe- 
cific conductivity, water transparency, substrate char- 
acteristics, and co-occurring fishes were recorded at 
all sampling locations except four that were devoid 
of cover and fishes. 

Fish were preserved in a 12:1 solution of 80% 
ethanol and glycerin, respectively, and stored at room 
temperature (22-25°C). A dissecting microscope 
equipped with an ocular micrometer was used for mea- 
surement and illustration of fish. Measurements <5 mm 
were accurate to +0.1 mm, and larger characters to 
+0.2 mm. Several specimens were cleared to determine 
vertebra number. Fish used in this study are stored at 
the Royal Ontario Museum, Toronto, Ontario. 

In general, terms used to describe fishes follow 
Trautman (1981). Base length of median fins is defined 
as the distance from the anterior margin of the first 
spine or soft ray to the posterior margin at the base of 
the last spine or ray. Snout length is the distance from 
the anterior margin of the premaxillary to the-anterior 
margin of the eye. Interorbital width is the least dis- 
tance between orbits. Body depth factor (BDF) and 
caudal depth factor (CDF) indicate general body 
form and swimming ability (Webb and Weihs 1986). 
Body depth factor is defined as mean TL/mean body 
depth, whereas CDF = mean caudal peduncle depth/ 
mean body depth. A high BDF and CDF denotes an 
elongate species whose depth is more or less the same 
along its length; e.g., clupeid or osmerid. Low values 
are found in centrarchids, where manoeuvrability is 
enhanced by a deep body and large pectoral fins. Fish 
volume, determined in each size class = mean total 
length x (mean body depth)’. 


Results 
Reproductive and nursery habitat 

Environmental conditions for fish reproduction and 
rearing differed in Duck Creek and the St. Clair River 
according to water quality and physical characteris- 
tics. On most dates, water temperature at the shore of 
the river was about 3°C lower and suspended particu- 
late load always lower, than in Duck Creek. Whether 
collected at 4° or 30°C, Round Gobies were recorded 
in the same habitat from April to November. However, 
their occurrence was contingent on availability of 
cover. For example, gobies were not found at LGS in 
May 1999 when low water level completely exposed 
rock and rubble habitat. Recently hatched fish were 
rarely found along featureless, sandy shores, where 


320 


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THE CANADIAN FIELD-NATURALIST 


Vol. 118 


SS ES 


eee a ae 
~ a] s 


FIGURE 2. Neogobius melanostomus at 7.5 mm; upper: dorsal, middle: lateral, lower: ventral view. Length bar = 1 mm. 


juvenile and adult gobies were occasionally caught. 
Small boat launches with corrugated concrete ramps 
were consistent sources of larger (>30 mm) specimens 
of age-0 gobies. 

Presence of small age-0 fish in late spring and early 
autumn indicates a possible prolonged spawning peri- 
od in Duck Creek, accommodated by water tempera- 
ture >10°C from early May to mid-October. Earliest 
evidence of recently hatched fish (5-13 mm TL) on 1 
June 1995 suggests spawning takes place in April or 
May. However, in early May 1998, ten gobies (22-28 
mm) were caught near LGS at 10°C above ambient 
(4°C). These fish may have hatched in late winter in 
dilute heated power plant effluent. Since small gobies 
(14-23 mm) were found each year in September, 
spawning at LGS may extend to late summer. 


In Duck Creek, Round Gobies endure continuous 
turbid water, enriched conditions, occasional dense 
blooms of blue-green algae (Anabeana sp; Cyanophy- 
ta) and temperature at least 30°C. According to size 
in autumn, gobies attain a mean length of about 33- 
39 mm at the end of first year growth. For example, 
in Duck Creek fish were 32.8 mm (28-36 mm) in early 
October 1995, whereas at LGS, they were 37.6 mm 
(range 31-49 mm) in November 1994 and 39.4 mm 
(30-45 mm) in November 1995. 


Reproduction 

In mid-July 1999, numerous clusters of eggs were 
found attached to the underside of rocks at a depth of 
0.3 m in Duck Creek. A sample (N = 121) of eggs in 
a cluster of roughly 200 on a rectangular patch of about 
30 cm? averaged 3.0 mm long and 1.8 mm wide with 


2004 


LESLIE and TIMMINS: AGE-O ROUND GoBY IN ONTARIO 32] 


FIGURE 3. Neogobius melanostomus at 25 mm. Length bar = 2 mm. 


respective modal values of 3.1 mm and 1.8 mm (range: 
2.6-3.3 mm and 1.6-2.0 mm). Eggs are oblong with 
rounded base and blunt apex, with a fibre-like basal 
pedestal or stalk 0.4-0.5 mm long. Yolk is light orange 
and occupies nearly 75% of the capsule. These eggs 
incubated in situ in turbid, almost lentic water at 28- 
30°C. Similarly, egg clusters were found in early June 
2000 at 14°C in the St. Clair River beneath limestone 
rocks and human artefacts. A cluster of approximately 
300 eggs formed an oval patch (estimated area = 
40 cm?) under a rock at a depth of 0.2-0.3 m in slow 
flowing water. Eggs (n = 78) were 3.5 mm long and 
2.0 mm wide (modal values, 3.7 mm and 2.1 mm). 


Morphology 

Yolk, which is usually retained in the gut of fish 
4.5 to 6.5 mm long, may persist to 11 mm. At 5-8 
mm, minute teeth are present on the jaws. Origin of 
D1 is at the fourth myomere or fifth vertebra, whereas 
D2 originates at the seventh or eighth myomere (elev- 
enth or twelfth vertebra). Scales first appear on mid- 
caudal peduncle at approximately 8-9 mm and body 
scalation may be complete at 12-15 mm. 

The upper lip is protractile and jaw articulation is 
below the anterior margin of the pupil. The anterior 
tubular nostril protrudes, whilst the minute posterior 


nostril is flush. At about 13 mm the cranium is de- 
pressed, cheeks begin to enlarge, and sub-orbital canals 
and papillae are barely visible. At all sizes, the head 
is bluntly arrow-shaped in ventral profile (Figure 2). 
All spines and fin rays are formed, although principal 
rays are incomplete in the caudal fin. Pectoral fin rays 
are joined distally by membrane. A small genital papilla 
is first obvious at about 23 mm. 

Origin of D2 is on a vertical line with the anus and 
separated from D1 by a flap of tissue. Full complement 
of branched caudal rays is attained at about 20 mm. 
At 22-25 mm, the pectoral fins are slightly flared ven- 
trally and depressed D2 and anal fins extend equally 
to mid-peduncle (Figure 3). Two short, stout spines 
support a transverse membrane on the pelvic fin, the 
whole forming a suctorial disc. 

Fish >30 mm have a moderately rounded snout 
and large protruding dorsolateral eyes above bulging 
cheeks. Sub-orbital papillae occur in transverse rows 
(Figure 4). The tip of the large fleshy upper lip is 
anterior to the lower, and the opening of the terminal 
mouth lies below the ventral margin of the eye. The 
centre of the pupil lies along a horizontal line with the 
origin of the pectoral fin. Gill openings are narrow. 
Ctenoid scales cover the body, whereas cycloid scales 
occupy the anterior nape and gill covers. In outline, 


Ficure 4. Lateral view of male Neogobius melanostomus at 38 mm. Length bar = 5 mm. 


B22 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE |. Morphometrics for age 0 Neogobius melanostomus collected in the St. Clair ecosystem, Ontario. Lengths are 
percentage (with range) of mean total length (TL) in each size class. 


Size 5.0-8.9 9.0-12.9 13.0-16.9 
class n= 20 m=9 n=2 
Mean TL Ussies A 10.4 + 1.2 13301 
Standard 79.0 79.3 78.1 
73-86 78-83 77-719 
Preanal 45.8 43.8 41.5 
43-52 42-45 41-42 
Predorsal 33.0 Silen 30.6 
30-42 30-33 30-32 
Prepelvic Pes A| PAE | 27.6 
18-30 24-31 27-28 
Body depth 17.8 16.8 1GS)59) 
14-21 15-18 14-17 
Peduncle US 8.0 8.3 
6-9 7-9 0 
Head 26.2 26.3 27.6 
21-30 24-29 27-28 
Eye? 30.6 Bilal 27.6 
25-37 26-35 27-28 
Volume (mm?*) 12.3 30.1 5i5).7/ 
BDF 5.6 6.1 6.5 
CDF 0.42 0.49 0.54 
Pectoral fin 18.6 19.7 D5 
10-22 17-22 21-22 
Pelvic fin 1525 Ne, 19.3 
9-21 13-19 18-20 
Dorsal | base 7.0 7.8 6.5 
5-11 6-10 6-7 
Dorsal 2 base 24.3 235 23.4 
19-30 20-29 23-24 
Anal base 20.5 19.7 18.5 
18-24 16-23 17-20 


*% head length 


D1 is slightly rounded, D2 slopes slightly, and the 
anal fin is uniform (Figure 4). 


Morphometry 

Relative changes in lengths are generally small as 
fish grow (Table 1). Head length (24-28% TL) is 1.3 
times its width and 1.4 times its depth. Snout length 
increased from 15% HL at 7 mm to 20% HL at 35 mm. 
Interorbital width, variable among and within size 
classes, is 10-22% HL. Pectoral fin rays are 19-24% 
TL whereas D2 base is 23-26% TL. Longest spines 
and base of D1 are about equal but less than the 
longest ray of D2. On fish >35 mm, the pectoral fin 
base is approximately 55% greatest body depth. Body 
depth and caudal depth factors are generally constant 
(Table 1). 
Meristics 

Respective modal counts for preanal, postanal, and 
total myomeres are 12 (range, 11-13), 19 (17-20), and 
31 (29-32). Modal spine and fin ray complements are 
D1: VI, D2: 1 15 (13-16), anal: I 13 (11-13), pectoral: 
18 (16-19), pelvic(s): 1 5 (conjoined), principal caudal: 


17.0-20.9 21.0-24.9 25.0-32.9 33.0-39.9 
n=9 n= n=33 N23, 
18.5+0.9 23.5+0.9 2837 = 21 35.3+ 1.8 
80.3 79.8 79.5 80.3 
79-83 78-83 77-81 79-82 
44.0 44.7 44.0 43.4 
43-46 41-48 42-51 39-46 
30.1 29.7 29.2 28.2 
29-32 28-32 27-31 24-29 
DAs 27.0 26.6 26.6 
26-28 24-32 25-29 25-34 
gh) 16.9 17.6 ei 
14-20 15-19 15-20 16-19 
9.0 8.9 8.9 8.8 
9-10 8-9 9-10 8-10 
25.5 253 24.8 24.0 
25-27 24-27 23-27 23-26 
33.0 32.4 B22 30.3 
31-35 28-35 28-35 26-35 
189 35 ii 2 1401 
5.8 6.0 Si 5.6 
0.53 0.52 0.51 0.50 
23.6 24.0 DES) 23.6 
22-25 22-25 19-27 21-26 
203 19.9 20.1 19.8 
20-22 18-21 18-22 17-23 
8.8 8.5 9.0 9.2 
8-9 7-12 8-11 7-11 
pip) 24.7 25.9 DS) 
24-28 20-29 24-27 24-27 
19.6 19.2 20.6 20.4 
17-22 17-22 17-23 19-23 


13 (11-13). There are 33 modal (32-33) vertebrae and 
49 (48-52) ctenoid scales in mid-lateral series. 


Pigmentation 

On free embryos, pigmentation consists of black 
eyes and several black spots on posterior spines of 
D1. However, on many specimens <9 mm, the char- 
acteristic black spot on D1 is absent. Speckled mel- 
anophores on snout and dorsum of head, a dark crescent 
on the occiput and a small patch on the cheek are 
typical at 10-12 mm. Ventrally, an elongate stellate 
melanophore is situated mid-gut. About five faint 
dorsolateral clusters are expressed on the body. There 
is a small group of stellate melanophores on frontal 
and parietal areas, a ventral patch on the pectoral fin 
base and a subcutaneous series on the intestine. Mel- 
anophores develop at the base of each anal fin ray, 
and a narrow mid-ventral series prevails from anal fin 
base to caudal fin base (Figure 2). At 14 mm, striation 
extends from maxilla to orbit and a black spot is 
pronounced between fifth and sixth spines on D1. 
Several faint mid-lateral blotches extend the length 
of the body. 


2004 


LESLIE and TIMMINS: AGE-O ROUND GoByY IN ONTARIO 323 


FiGuRE 5. Cottus bairdi at 10.5 mm. Length bar = | mm. 


At 22-25 mm a large brown patch is obvious on 
the dorsal peduncle of the pectoral fin. First four rays 
of D1 are lightly pigmented, and patches appear on 
the base of interradials. A faint stripe exists mid-fin on 
D1 (Figure 3). Posterior margins of body scales are 
darkly outlined and rays at the base of the caudal fin 
are lightly pigmented. A small basicaudal spot persists 
throughout development. 

Fish >30 mm have a small stripe on the preopercle, 
and about seven irregular patches of brown pigment 
on the side of the body, mainly below the mid-line. 
Otherwise, lips and dorsum of head are covered with 
small round spots, whilst the throat is essentially de- 
void of pigment. Pelvic and caudal fins are lightly 
spotted, whereas D2 has two pale longitudinal stripes. 
Pigmentation patterns are intensified over head, body, 
and fins (Figure 4). Overall colouration on individuals 
varies from dark brown to grey-brown. 


Description of age-O Mottled Sculpin 

Mottled Sculpin larvae (11.4 + 1.1 mm; n = 5) 
have the following characters: preanal length 41% 
TL, head 25%, eye 29% head length; head length and 
width equal; predorsal length 29% TL, prepelvic length 
24%. Branchiostegals 3,4. Except for pelvic fin sep- 
aration, overall body characters are similar to gobies 
(Figure 5). Pigmentation features black eyes and 
approximately six lateral saddle marks on the body. 
Melanophores are absent on the ventral body surface 
and fins. Meristics for this, and superficially similar, 
species are given in Table 2. 


Discussion 

The Round Goby is an eurythermal fish adapted to 
slightly brackish environments in the Caspian and 
Black seas (Miller 1986). In the Great Lakes basin it 
is ubiquitous in the littoral zone, including ecotones 
intolerable to many of our native fishes. Eggs incu- 
bate in almost static water in Duck Creek, a stream 
burdened with silt, and at times, blooms of blue- 
green algae at high temperatures. Concurrently, eggs 
may develop in the St. Clair River in water of high 
quality and clarity at temperatures several degrees 
lower than in small turbid streams in the same catch- 
ment. The reproductive strategy includes parental 
guarding of eggs and free embryos. In the St. Clair 
aquatic ecosystem, age-O0 Round Gobies appear to 
remain natal stream and shore habitats throughout 
their first year of growth. Solid objects, unlike sand 
and other unstable substrates, provide surfaces essential 
for attachment and refuge for small gobies in flowing 
water. 

Reproduction is prolonged and extensive but varies 
geographically. In their native range, gobies migrate 
to spawning habitat at 9-26°C and reproduce from 
April-September in intervals of 3 or 4 weeks (Charle- 
bois et al. 1997). Similar data for the Great Lakes are 
lacking, although MacInnis and Corkum (2000) suggest 
a possible spawning period from April to November 
in the upper Detroit River. In power plant cooling 
water discharges, the reproductive period may extend 
from winter to late summer or beyond, since the endo- 
crine system is active all year (Charlebois et al. 1997) 
and water temperatures are constantly >10°C. 


TABLE 2. Comparison of meristics of several Great Lakes age-0 fishes superficially similar in appearance. 


Species D1 D2 Pelvic Pectoral Anal Preanal _Postanal 
fin fin fin fin myomeres myomeres References 

Etheostoma exile VII-X LO=12 5 12-14 II, 7-8 Heufelder 1982 
Etheostoma nigrum VIl-XI 10-14 5 11-12 EWVealO els Bil Heufelder 1982 
Etheostoma caeruleum x 12 6 13 ind) 16-17 18-19 Cooper 1979 
Cottus bairdi VIL-IX 16-19 I, 3-4 13-17 13-14 9-12 19-21 Heufelder 1982 

VIL-VIN =15-17_ I, 3-4 13-15 13 9-10 19-21 This study 
Cottus cognatus VII-IX 16-19 1, 3-4 12-16 10-14 9-12 19 Heufelder 1982 
Proterorhinus marmoratus VI-VII 16-19 1,5 14-15 | is) 11 20 Leslie et al. 2002 
Neogobius melanostomus VI 1S =1G) les 18 iatles} 12 19 This study 


324 


In the study area, batches of about 200-300 eggs 
represent an absolute fecundity of 800-1200, assuming 
four spawning episodes. MacInnis and Corkum (2000) 
estimated an absolute fecundity of 252-1818 eggs based 
on three spawning episodes from late May to the end 
of July in the upper Detroit River. Miller (1986) report- 
ed fecundity of gobies 7-13 cm long as 328-5221 in the 
Black-Caspian-Azov seas, whilst Kuczyfiski (1995) 
calculated 2700-3000 eggs in the Gulf of Gdansk. In 
general, fecundity is lower and eggs slightly smaller 
(3.7-3.9 mm long, 2.0-2.2 mm wide) than in the Black 
and Baltic seas, possibly because the spawning popu- 
lation is younger and fish are smaller (MacInnis and 
Corkum 2000). 

Moskal’kova (1967) reared embryos that emerged 
from the egg at 5.5-5.7 mm and remained in the nest 
4-9 days. Gobies do not have a true larval stage (Miller 
1986) and hatch with most characteristics of juveniles. 
However, in the present study, smallest gobies (4.5- 
5.0 mm) feature a large yolk sac and incomplete com- 
plement of fin rays. These fish may have extruded from 
eggs during the collection process and perhaps are not 
representative of free embryos. Otherwise, smallest 
specimens found in coarse gravel were clearly un- 
guarded, free-living fish. 

The Round Goby is a cryptic, benthic fish, devoid 
of air bladder and planktonic stage. As such, early 
developmental stages are found in and among firm 
structures. Gobies have smaller caudal and body depth 
factors than larvae capable of high thrust and acceler- 
ation, such as Anchovy (Eugraulidae), Alewife (Clu- 
peidae) or percids (C. A. Timmins, unpublished, 2000). 
The Round Goby may not be a strong or fast swimmer, 
but its large head, robust body, and large pectoral fins 
enhance its manoeuvrability. 

Age-0 Round and Tubenose gobies did not co-occur 
in any given habitat in the St. Clair ecosystem, al- 
though they partitioned habitat in a small area near 
the mouth of Duck Creek (Leslie and Timmins 1998). 
Usually, small age-O Round and Tubenose gobies may 
be separated on the basis of head and body pigmen- 
tation, which is much lighter in the former. Whereas 
the Round Goby has a conspicuous black spot on the 
posterior spines of D1, the Tubenose Goby has an 
oblique stripe on anterior spines. Further, tubular nos- 
trils that extend beyond the upper lip are peculiar to 
the Tubenose Goby. However, most mensural char- 
acters overlap for these gobies. Thus, small (<6 mm) 
unpigmented fish may not be separable. 

Fishes with similar habitat requirements and mor- 
phology, such as cottids, darters, and gobies sometimes 
co-occur. Recently hatched Mottled Sculpins are pre- 
cocious and appear similar to gobies (Figure 5). 
However, all fins are developed in recently hatched 
Round Gobies, whereas cottids (and darters) have a 
complete fin fold and separate pelvic fins, which are 
expressed some weeks post-hatch. In addition, pre- 
anal and total myomere counts of cottids and darters 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


do not overlap with those of gobies. Fin spine and ray 
counts and pigmentation patterns also differ among 
these taxa. 

The Round Goby has survival advantage over many 
competitors. It reproduces repeatedly from spring to 
autumn, the critical life stage is not determined by 
specific seasonal type or quantity of food, its embryos 
hatch with almost all external structures elaborated, 
the young are guarded in nests and it persists in natal 
habitats. Certainly, few native fishes of the Great Lakes 
basin are as morphologically and physiologically suit- 
ed to thrive in comparable environmental conditions. 
Therefore, geographic range expansion and prolifer- 
ation of the species seems likely in the Great Lakes 
Basin. 


Acknowledgments 

R. E. Dermott, Department of Fisheries and Oceans, 
Burlington, Ontario, kindly provided specimens of 
mottled sculpin. We thank two anonymous reviewers 
for comments that improved the manuscript. 


Documents Cited (marked * in text) 

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Research Network on ecology of marine invasions and 
introductions. Edited by S. Olenin and D. Daunys. Internet: 
http: www.ku.lt/nemo/mainmeno.htm. 


Literature Cited 

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Wolfe, D. J. Jude, and S. Rudnicka. 1997. The round 
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pean and North American literature. Ilinois-Indiana Sea 
Grant Program and Illinois Natural History Survey. 
Special Publication Number 20. 

Cooper, J. E. 1979. Description of eggs and larvae of Fan- 
tail (Etheostoma flabellum) and Rainbow (E. caeruleum) 
darters from Lake Erie tributaries. Transactions of Amer- 
ican Fisheries Society 108: 446-456. 

Crossman, E. J., E. Holm, R. Cholmondeley, and K. 
Tuininga. 1992. First record for Canada of the Rudd, 
Scardinius erythrophthalmus, and notes on the introduced 
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Dubbs, D. O. L., and L. D. Corkum. 1996. Behavioural 
interactions between Round Gobies (Neogobius melanos- 
tomus) and Mottled Sculpin (Cottus bairdi). Journal of 
Great Lakes Research 22: 838-844. 

Heufelder, G. R. 1982. Family Cottidae, sculpins. Identi- 
fication of larval fishes of the Great Lakes basin with 
emphasis on the Lake Michigan drainage. Pages 1-774 in 
Great Lakes Fisheries Commission Special Publication 82- 
83. Edited by N. A. Auer. Ann Arbor, Michigan. 

Jude, D. J., R. H. Reider, and G. R. Smith. 1992. Estab- 
lishment of Gobtidae in the Great Lakes Basin. Canadian 
Journal of Fisheries and Aquatic Sciences 49: 416-421. 

Kalinina, E. M. 1976. [Abstract] Reproduction and early 
development of gobies from the Azov and Black seas. 
Kiev. Nauk Dumka, Russia. 

Koblickaja, A. F. 1981. [Abstract]. Opredielitiel molodii 
priesnowodnyh rby Moskva. Legkaja i pischevaja pro- 
myslennost 208. [cited in Charlebois et al. 1997]. 


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Kuczynski, J. 1995. Babka kragtia Neogobius melanostumus 
(Pallas 1811) — emigrant z Basenu Pontokaspijskiego w 
zatoce Gdanskiej. Bulletin Sea Fisheries Institute, Gdynia 
2(135): 68-71. 

Leslie, J. K., and C. A. Timmins. 1998. Fish reproduction 
and distribution in a small tributary of Lake St. Clair. Can- 
adian Technical Report of Fisheries and Aquatic Sciences 
2253): 

Leslie, J. K., C. A. Timmins, and R. G. Bonnell. 2002. 
Postembryonic development of the tubenose goby Protero- 
rhinus marmoratus Pallas (Gobiidae) in Lake St. Clair 
and the St. Clair River. Archivum fiir Hydrobiologie 154: 
341-352. 

MacInnis, A. J., and L. D. Corkum. 2000. Fecundity and 
reproductive season of the Round Goby Neogobius melan- 
ostomus in the upper Detroit River. Transaction of 
American Fisheries Society 129: 136-144. 

Miller, P. J. 1986. Gobiidae. Pages 1019-1095 in Fishes of 
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head, M.-L. Bauchot, J.-C. Hureau, J. Nielsen, E. Torto- 
nese. Volume 3: UNESCO, Paris. 

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development of the Round Goby, Neogobius melanostomus 
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Received 17 February 2003 
Accepted 15 November 2004 


A Systematic Analysis of the Alpine Saxifrage Complex (Saxifragaceae) 
in the Canadian Arctic Islands Using Morphology and Chloroplast 
DNA Data 


CAROLINE HEALY! and LYNN J. GILLESPIE 


Research Division, Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4, Canada 
' Present address: Department of Biology, University of Ottawa, P.O. Box 450, Station A, Ottawa, Ontario KIN 6N5 Canada 


Healy, Caroline, and Lynn J. Gillespie. 2004. A systematic analysis of the alpine saxifrage complex (Saxifragaceae) in the 
Canadian Arctic Islands using morphology and chloroplast DNA data. Canadian Field Naturalist 118(3): 326-340. 


The Saxifraga nivalis complex displays significant ecological, morphological and cytological variation. Most European studies 
suggest that the S$. nivalis complex comprises two distinct species: Saxifraga nivalis sensu stricto and Saxifraga tenuis. 
However, the presence of intermediate morphotypes, inconsistencies in chromosomal counts and variability in morphological 
keys and descriptions have led to different taxonomic interpretations of the complex in North America. This study investi- 
gated the systematics of Canadian Arctic Island members of this complex from 157 specimens using 23 morphological 
characters. Principal component analysis of the morphological data revealed two adjacent clusters, corresponding to the two 
taxa and consistent with a close morphological similarity and the presence of hybrids. A preliminary restriction site analysis 
of five non-coding regions of the chloroplast genome, trnH-trnK, trnT-trnF, trnF-trnV, trnV-rbcL and rbcL-ORF106, was 
conducted using 21 restriction endonucleases. This analysis indicated a length difference between the trnT-tvnF region of S. 
nivalis and that of S. tenuis, but no difference in restriction sites for any of the assayed regions. These results confirm that in the 
Canadian Arctic, the S. nivalis complex consists of two closely related, largely sympatric species, with notable morphological 
variability, and possible hybrids. 


Key Words: Alpine Saxifrage, Saxifraga nivalis, Saxifraga tenuis, Canadian Arctic, systematics, morphology, chloroplast 


DNA restriction site analysis. 


Saxifraga L., the largest genus in the Saxifragaceae, 
consists of nearly 400 herbaceous, mostly perennial 
species (Webb 1993; Soltis et al. 1993, 1996). Saxi- 
fraga nivalis L. sensu lato, commonly known as the 
Alpine Saxifrage, inhabits arctic and alpine environ- 
ments throughout the northern hemisphere. The S. 
nivalis complex is dispersed throughout northern and 
alpine Europe, Siberia, arctic and subarctic North 
America and south in the alpine zone to Arizona and 
the Gaspé, Quebec (Britton and Brown 1913; Krause 
and Beamish 1973; Scoggan 1978; Webb 1993). 
Members of the S. nivalis complex display notable 
morphological, cytological and ecological variation, 
which has led to different taxonomic interpretations 
and classifications. 

Linnaeus (1753) first described S. nivalis, a name 
that means snow saxifrage, as these plants are often 
found growing in snow patch plant communities (Brit- 
ton and Brown 1913). Later, Wahlenberg (1812) recog- 
nized S. nivalis var. tenuis Wahlenb. for plants with a 
notably smaller stature, fewer flowers, smaller fruits 
and recurved stigmas. Smith (in Lindman 1918) elevat- 
ed Wahlenberg’s variety to Saxifraga tenuis (Wahlenb.) 
Harry Sm. However, the taxonomy of the S. nivalis 
complex remained problematic and the acceptance of 
S. tenuis as a species was controversial. For instance, 
Bocher (1938) recognized S. nivalis and S. tenuis as 
distinct species, but also suspected the presence of races 
and microspecies within the S. nivalis complex. Polunin 
(1940) found the two taxa difficult to separate morpho- 


logically with intermediates and suggested the need for 
breeding and cytological studies before distinguishing 
them at the species level. Although accepted as a dis- 
tinct species in Scandinavia and Greenland, based on 
samples from Alaska and the Yukon, Hultén (1945) 
concurred at the time with Polunin in treating the 
taxon as S. nivalis var. tenuis because of its minimal 
phenotypic distinction from S. nivalis and its low fre- 
quency of occurrence within the area of distribution of 
S. nivalis. However, in the Canadian Arctic, Porsild 
(1957, 1964) and Porsild and Cody (1980) recognized 
S. nivalis and S. tenuis as distinct species. Upon fur- 
ther examination of specimens from Alaska and neigh- 
boring regions, Hultén (1968) also recognized the 
two taxa at the species level. More recently Scoggan 
(1978) and Aiken et al. (1998) treat the two taxa as 
varieties of S. nivalis. 

Taxonomic and nomenclatural ambiguity also 
stemmed from the description of additional varieties and 
microspecies, as well as the re-emergence of Haworth’s 
(1812) suggestion to revise Linnaeus’ concept of the 
genus Saxifraga (reviewed in Spongberg 1972). In 
1905, Small divided the genus Saxifraga into 13 genera 
in North America (in Small and Rydberg 1905; 
reviewed in Soltis et al. 1996). Small treated S. nivalis 
as Micranthes nivalis (L.) Small, S. nivalis var. tenuis 
as Micranthes tenuis (Wahlenb.) Small, and distin- 
guished an additional species in the Canadian eastern 
Arctic, Micranthes kumlienii Small. However, subse- 
quent authors did not recognize the genus Micranthes, 


326 


2004 


and M. kumlienii was treated as belonging to S. nival- 
is (Polunin 1940; Aiken et al. 2000*). Fernald (1917) 
distinguished Saxifraga nivalis var. labradorica Fern., 
a variety localized in Labrador, and suggested that this 
plant corresponded to Small’s concept of Micranthes 
tenuis. Subsequently, this variety was treated as a syn- 
onym of S. nivalis var. tenuis (Scoggan 1978) or as a 
luxuriant form of the latter (Polunin 1940). In the Gaspé 
Peninsula, Fernald (1917, 1950) identified another 
member of the S. nivalis complex, Saxifraga gaspensis 
Fern. Subsequently, this species was included under 
S. nivalis var. tenuis (Scoggan 1950, 1978), treated as 
S. nivalis var. gaspensis (Fern.) Boivin (Boivin 1966), 
and then considered as small S. nivalis s.s. (Blondeau 
1989a). Most recently, Gervais et al. (1995) followed 
Fernald in treating the Gaspé population as a distinct 
species. In Alaska, Hultén (1968) considered S. nivalis 
as highly variable, recognizing S. nivalis var. tenuis, 
and describing S. nivalis var. rufopilosa Hultén for 
plants characterized by “densely pubescent, rufous- 
haired leaves and red petals” (page 579) found in 
Alaska and the Yukon. Porsild (1957, map page 188) 
appeared to treat S. rufopilosa under S. tenuis ac- 
cording to his map of that species (see also Porsild and 
Cody 1980, map page 403; Cody 1996). Later, Porsild 
(1975) recognized the latter as the distinct species, S. 
rufopilosa (Hult.) A. E. Porsild, and suggested that it 
was more closely related to S. tenuis than S. nivalis. 
Cody (1996) also recognized S. rufopilosa as a distinct 
species and considered S. tenuis as not present in the 
Yukon. In 1978, Scoggan’s concept of S. nivalis var. 
tenuis encompassed S. tenuis, S. gaspensis, S. nivalis 
var. labradorica and S. nivalis var. rufopilosa. 

Based on cytology and morphology, Love (1983) 
and Webb (1964, 1993) discerned the species S. nivalis 
(2n = 60) and S. tenuis (2n = 20) in Europe. The taxo- 
nomic ambiguity regarding the S. nivalis complex in 
North America (Aiken et al. 2000*) appears to stem 
from the following factors: (1) several morphological 
characters used to distinguish the species in Europe 
do not hold up in North America; (2) taxa can be found 
growing together; and (3) intermediate morphotypes, 
as well as variation in chromosomal counts suggest 
hybridization. Various cytological studies of material 
from North America have demonstrated cytological 
diversity within the S. nivalis complex with chromo- 
somal counts of 2n = 20, 40, c. 56, 58, 60 or c. 60. 
While most support the above European counts for S. 
nivalis and S. tenuis (Mosquin and Hayley 1966; Hed- 
berg 1967; Johnson and Packer 1968; Packer and 
McPherson 1974; Love and Love 1975), several studies 
found intermediate chromosome counts. Krause and 
Beamish (1973) provide counts for plants of the S. 
nivalis-tenuis complex from the Yukon (2n = 20, 40), 
British Columbia (60) and Idaho (20), which they were 
unable to identify to species. Intermediate chromosome 
counts have also been found in the Russian Arctic in 
a plant described as S. nivalis x S. tenuis (Devyatov et 


HEALY AND GILLESPIE: ALPINE SAXIFRAGE COMPLEX 327 


al. 1997). In the Gaspé Peninsula S. gaspensis is con- 
sidered to be a putative endemic hybrid based on chro- 
mosome number (2n = 40) and apparent intermediate 
morphology (Gervais et al. 1995). Further confusion 
results from the variance and discrepancies among the 
morphological keys used in the identification of this 
complex and related species, such as S. foliolosa R. 
Br., which are occasionally misidentified as members 
of the S. nivalis complex. Consequently, accurate and 
consistent identification of North American plants 
belonging to this complex can be problematic. 

The aim of this study was to investigate the distinc- 
tiveness of S. nivalis and S. tenuis in the Canadian 
Arctic Islands based on morphological and molecular 
differences, and to help reassess the taxonomic status 
of S. tenuis. 


Material and Methods 
Morphological Analysis 
Specimens 

Representative specimens from the Canadian Arc- 
tic were selected from the Canadian National Herb- 
arium, Canadian Museum of Nature (CAN), and the 
Agriculture and Agri-foods Canada Herbarium, Cen- 
tral Experimental Farm, Ottawa (DAO). Morphological 
data were gathered from observations and measure- 
ments on 157 herbarium sheets (listed in Appendix 
I), including two chromosome count vouchers (S. 
nivalis, 2n = 60, Calder et al. DAO 24148: S.. tenuis, 
2n = 20, Calder et al. DAO 24149). One plant per 
sheet was sampled, except when sheets comprised 
mixed collections or included DNA vouchers, and 
then multiple plants per sheet were sampled. A total 
of 200 plants were sampled, 122 S. nivalis and 78 S. 
tenuis. More specimens of S. nivalis were available for 
study than of S. tenuis. Where identifications of exam- 
ined specimens were judged questionable, specimens 
were re-identified based on a combination of published 
keys and preliminary results. Additionally, identifica- 
tions of many S. tenuis specimens at CAN were sub- 
sequently confirmed for the Panarctic Flora Project 
(E. Reidar, 2001, 2002). 


Morphological characters 

Characters used in descriptions and keys of the S. 
nivalis complex (Simmons 1906; Polunin 1940; Porsild 
1957; Love 1983; Blondeau 1989a, 1989b; Webb 
1993; Aiken et al. 1998, 2000*) were reduced to a set 
of 30 characters based on ease of observation or meas- 
urement on dried specimens as well as their taxonom- 
ic significance. Of these, 23 were selected for statis- 
tical analysis (Table 1). Petal length was excluded 
from all analyses due to difficulties in measuring petals 
that were often folded and partly included within the 
calyx. The qualitative characters, stem colour, petal 
shape, prominence of leafy bract on stem, leaf blade 
shape, leaf blade base shape and apex shape, were 
excluded due to variation within a plant or difficulties 
in determining discrete states. 


328 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE |. Morphological characters used in analyses of the Saxifraga nivalis complex in the Canadian Arctic. 


Quantitative characters: 


Hair length on leaf lower surface (mm): UNDHRLGTH 
Hair length along leaf margin (mm): MRGHRLGTH 


SOC eE ES ON Calipers) Ei 


Plant height (cm): HEIGHT; length from basal rosette to top of inflorescence 

Stem width (mm): STWIDTH; measured midway along lower half of stem 

Stem hair density (0, glabrous to 4, dense): STHRDENSITY 

Stem hair length (mm): STHRLENGTH; measured midway along lower half of stem. 
Leaf blade length (cm): LFLENGTH; measurement made on the largest leaf 

Leaf blade width (em): LFWIDTH; measurement made on the largest leaf 


Number of teeth on leaf: TEETH; count made on the largest leaf 


10. Tooth width (mm): TTHWIDTH; measurement made on largest tooth on the largest leaf 
11. Tooth length (mm): TTHLENGTH; measurement made on largest tooth on the largest leaf 
12. Petal width (mm): PTWIDTH; measurement made on the largest petal 


Qualitative characters: 

Stem hair texture (short fine; long coarse) 
Tooth apex shape (rounded; pointed) 

Hair colour on leaf lower surface (rust; white) 
Hair colour on leaf margin (rust; white) 


Inflorescence density (open; tight cluster) 
10. Style shape (curved; straight or essentially so) 


SEN SION a eS 


11. Petal colour (pink or essentially so; white or essentially so) 


The software package DELTA (Dallwitz et al. 1993) 
and the character set for the “Flora of the Canadian 
Arctic Archipelago” (Aiken et al. 2002*) were used to 
generate preliminary taxon descriptions. These descrip- 
tions were then modified to read more easily, to ex- 
clude irrelevant characters, and to include important 
diagnostic characters not included in the Flora char- 
acter set. 


Statistical analysis 

All statistical analyses were performed with SYSTAT 
version 8.0. For each quantitative trait, the means for 
S. tenuis and S. nivalis specimens were compared by 
performing t-tests. Separate variance values were cal- 
culated for each taxon. T-tests were conducted on both 
raw and log-transformed data. A principal component 
analysis (PCA) using the 12 quantitative characters in- 
dicated in Table 1 was conducted to visualize possible 
distinctiveness between S. nivalis and S. tenuis. The 
character MRGHLGTH was excluded from some PCA 
analyses due to a large amount of missing data in S. 
tenuis, which sometimes lacked marginal hairs. The 
PCA analysis excluded all specimens with missing 
values. A discriminant function analysis was also per- 
formed on the quantitative data. For the 11 selected 
qualitative characters (Table 1) the percentages of spec- 
imens having each character state was determined for 
each taxon. 

To determine if there was any geographical pattern 
to the morphological variation in S. tenuis, principal 
component analysis was performed only on the S. 
tenuis quantitative data (excluding MRGHLGTH). 
This analysis also included five specimens from Alas- 


Stem hair colour (rust-coloured; mixed rust and white; white) 


Regularity of teeth (even in size; variable in size smaller towards leaf blade apex) 
Hair presence on leaf lower surface (moderate to dense; sparse; absent) 


Hair presence on leaf margin (moderate to dense; sparse; absent) 


ka (Calder 6107, 6511, 6249 DAO) and northern 
British Colombia (Calder & Kukkonen 28151 DAO, 
Taylor et al. 1204 DAO) not included in the larger 
analysis. Individuals were coded as occurring in one 
of three regions: high arctic (67), eastern arctic (11) 
or western montane (5). 


Chloroplast DNA analysis 
Plant material and DNA extraction 

A total of 15 plants, for which silica gel dried mate- 
rial was available, were selected for molecular analysis: 
seven individuals of S. nivalis (Gillespie 6165, 6675-1, 
6826a-1, 6826a-2, 6977-2, 6984-5; Aiken 98-055-3), 
seven S. tenuis (Gillespie 6728-1, 6807-1, 6807-3, 
6825-2, 6870-2, 6877-1, 6881-3), and one S. foliolosa 
(Gillespie 6841) (Appendix 1). The latter species was 
selected to provide perspective on the chloroplast 
DNA (cpDNA) data, based on its close phylogenetic 
relationship to the S. nivalis complex (Soltis et al. 
1996). DNA was extracted following Doyle and Doyle’s 
(1990) CTAB total DNA extraction procedure as 
modified by Gillespie et al. (1997). 


Restriction site analysis 

DNA extracts were PCR amplified using five pairs 
of universal primers for non-coding regions of chloro- 
plast DNA. The five regions are: tmmH-tmmK (Demesure 
et al. 1995), trnT-trnF (Taberlet et al. 1991), trnF-trnV 
(Dumoulin-Lapegue et al. 1997), trnV-rbcL (Dumou- 
lin-Lapegue et al. 1997), and rbcL-ORF106 (Arnold 
et al. 1991). Amplification reaction mix and programs 
follow Gillespie and Boles (2001). 


2004 HEALY AND GILLESPIE: ALPINE SAXIFRAGE COMPLEX 329 


TABLE 2. Descriptive statistics and t-test comparing the means of S. nivalis and S. tenuis at the 95% confidence level for 12 
quantitative morphological characters. 


Range Difference 
Character Taxon N Mean (+SD) Min Max in means t value df p 


S. nivalis 122 OFZ) CE3'3i7) af AsO) 


Plant height (cm) 4.66 13.68 176.2 0.000 
S. tenuis 78 4.40 (+1.40) 1.8 8.5 
S. nivalis 122 1.37 (+0.35) 0.7 DES 

Stem width (mm) 0.82 23.06 178.6 0.000 
S. tenuis 78 0.56 (+0.15) 0.3 1.0 
S. nivalis 122 3.2 (+ 0.70) 1 4 

Stem hair density 2.0 24.94 205.0 0.000 
S. tenuis 78 1.2 (+ 0.36) 1 3} 
S. nivalis 1:22 3.2 (+ 0.70) 1 4 

Stem hair length (mm) 0.67 35.45 188.5 0.000 
S. tenuis Tal 0.19 (+0.09) 0.1 0.5 
S. nivalis 119 0.96 (+0.34) 0.4 Dil 

Leaf blade width (cm) 0.51 14.81 161.2 0.000 
S. tenuis 77-044 (+0.13) 0:25 570!8 
S. nivalis 119 1.38 (+ 0.50) 0.6 3.6 

Leaf blade length (cm) 0.69 13.51 166.5 0.000 
S. tenuis TL 0.67 (+ 0.18) 0.4 IES 

Hair length on lower S. nivalis 120 0.38 (+0.13) 0.2 12, 

leaf surface (mm) 0.10 725) 97:08 70/000 
S. tenuis 78 0.27 (+0.06) 0.1 0.5 

Hair length on S. nivalis 121 0.19 (+ 0.07) 0.1 0.4 

leaf margin (mm) 0.07 8.47 167.4 0.000 
S. tenuis 55% 0.12 (+ 0.04) 0.1 0.2 

Number of teeth S. nivalis 118 10.3 (+1.74) 7 14 

on leaf blade 2.8 16.63 167.8 0.000 
S. tenuis 78 7.5 (40.73) 6 9 
S. nivalis 118 1.17 (+ 0.42) 0.5 3h) 

Tooth width (mm) 0.37 7.86 195.6 0.000 
S. tenuis 78 0.80 (+ 0.24) 0.5 1.6 
S. nivalis 118 1.52 (+0.48) 0.8 B82. 

Tooth length (mm) 0.62 11.84 190.5 0.000 
S. tenuis 78 0.90 (+0.26) 0.4 1.8 
S. nivalis 120 1.12 (+0.24) 0.7 1.8 

Petal width (mm) 0.40 14.16 202.0 0.000 
S. tenuis al 0.72 (+0.16) 0.5 il 


* Marginal hair was not present in all S. tenuis 


PCR products were single digested with twenty one Apal, BamHI, Bgil, BglII, Dral, EcoRI, EcoRV, Hincll, 
restriction endonucleases according to manufacturer’s PstI, XhoI. To separate restriction fragments, 8 uL of 
recommendations. These included nine 4-base pair the reaction mixture and 2 uL of buffer-dye mix were 
(bp) cutting restriction enzymes: Alul, Haelll, Hhal, run on 1.1% agarose gels with ethidium bromide at 
Mspl, Rsal, Ddel, Hinfl, Sau96], Taql; two 5bp cutting 38 V for 2-3hrs. The restriction fragment patterns were 
restriction enzymes: BstOl, SinI; and ten 6bp enzymes: visualized and photographed under ultraviolet light. 


330 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE 3. Distribution of qualitative morphological character states in Saxifraga nivalis and S. tenuis in the Canadian Arctic. 
Numbers represent percentage of individuals having a particular character state. 


Character Character state 
Stem hair colour rust 
mixed 
white 
Stem hair texture short fine 
long coarse 
Tooth apex shape rounded 
pointed 
Regularity of teeth even-sized 
variable 
Hairs presence on leaf lower surface moderate-dense 
sparse 
absent 
Hair colour on leaf lower surface rust 
white 
Hair presence on leaf margin moderate-dense 
sparse 
absent 
Hair colour on leaf margin rust 
white 
Inflorescence density open 
tight cluster 
Style shape curved 
straight 
Petal colour pink 
white 


Lengths of total amplified product and restriction frag- 
ments were estimated by comparison with known 
100 bp ladder DNA markers. 


Results 
Morphological Analysis 

To determine whether the means of the examined 
quantitative characters were different between S. nivalis 
and S. tenuis, t-tests were performed (Table 2). The 
analysis revealed a significant difference in means be- 
tween the two taxa for all characters examined. Overall, 
plants belonging to S. nivalis are more robust than S. 
tenuis as implied by higher means in both plant height 
and stem width observed in S. nivalis. Similarly, leaves 
of S. nivalis are on average longer, wider and have 
longer, wider and more numerous marginal teeth than 
S. tenuis. Also, hairs on the stems are longer and denser 
and hairs on the lower surface and margin of leaves 
are on average longer in S. nivalis. Petal width was sig- 
nificantly greater in S. nivalis than in S. tenuis. How- 
ever, the two taxa have notably different variances, 
with S. nivalis having a higher variance than S. tenuis 
for all quantitative characters examined. Considering 
the very low probability values and the robustness of 
t-tests for samples of this size, the differences in means 
can be considered as meaningful. Analyses were also 
performed on log-transformed data, with no significant 
differences. 

The frequency of occurrence of states of qualitative 
characters differed considerably between the two taxa 


S. nivalis S. tenuis 
3} 74.4 
0 25.6 

98.7 0 
1.6 100 
98.4 0 
DNS 100 
97.5 0 
18.3 87.2 
81.7 12.8 
91.7 84.6 
es 15.4 
0.8 0 
40.0 79.5 
60.0 20.5 
91.7 10.3 
8.3 60.3 
0 29.5 
20.7 60.0 
79.3 40.0 
1.8 97.1 
98.2 2.9 
25.9 84.7 
74.1 15}3) 
14.9 81.8 
85.1 18.2 


for most characters (Table 3). The characters, stem hair 
colour and texture, tooth apex shape and inflorescence 
density, differed significantly between the two taxa, 
with less than a five percent overlap in states. Several 
characters, regularity of teeth, petal colour and style 
shape, exhibited an overlap in states of 10-26% between 
the two taxa. In contrast, one character, hair presence 
on lower leaf surface, displayed very similar frequen- 
cies of states between S. nivalis and S. tenuis, with 
hairs present in over 95% of the specimens examined 
in both taxa, although density was sometimes more 
sparse in S. tenuis. The remaining characters displayed 
considerable overlap in states between the taxa. 

The principal component analysis of quantitative 
characters (excluding MRGHLGTH) revealed two 
clusters, positioned adjacent to each other with little 
overlap (Figure 1). Factor loading for the analysis was 
STWIDTH, LFWIDTH, LFLENGTH, STHRLENGTH, 
STHRDENSITY, HEIGHT, TEETH, PIWIDTH and 
TTHLENGTH for factor 1, and TTHWIDTH and 
UNDHRLGTH for factor 2, with 61% and 10% of total 
variance explained by factors | and 2, respectively. As 
noted above, there is considerable variation among S. 
nivalis plants for most examined characters. This can 
be observed on the graph through the large oblique 
spread of S. nivalis compared to S. tenuis. The PCA 
including the character MRGHLGTH produced very 
similar results (although displayed fewer individuals). 
The discriminant function analysis performed on the 
same data distinguished between the two taxa 100% 


2004 

5 

4 

3 

2) ORO 
Q 1 On S Boo GS ‘OP 
2 © 
O 
<= 
LL 

9) 

ES 

-4 

5 

5 -1 0 { : : 

FACTOR(1) 


FIGURE |. Principal component analysis on the quantitative 
morphological data set for the Saxifraga nivalis com- 
plex in the Canadian Arctic. Circles represent individ- 
uals of S. nivalis, crosses represent S. tenuis. 


of the time, with one dimension accounting for the 
variation. 

The PCA analysis examining geographical patterns 
to morphological variation in S. tenuis revealed very 
little geographical structure, with eastern arctic and 
high arctic plants almost completely overlapping. 


HEALY AND GILLESPIE: ALPINE SAXIFRAGE COMPLEX 33] 


Western montane plants separated from the majority 
of arctic plants on a combination of both factors (with 
LFWIDTH, LFLENGTH, and TEETH, loading high 
on factor one, HEIGHT on factor two), but overlapped 
considerably with several eastern arctic plants. A dis- 
criminant function analysis performed on the three 
geographic groups produced similar results (correctly 
placing the western montane plants with 100% reli- 
ability, high arctic plants with 86% reliability, and 
eastern arctic plants with only 42% reliability). 


Chloroplast DNA analysis 

The trnH-trnK amplicons were estimated at 1600 
base pairs (bp) for S. nivalis and S. tenuis and 1700 bp 
for S. foliolosa. The three largest regions, trnF-trnV, 
trnV-rbcL and rbcL-ORF106, were approximately 
3000 bp in length in all three taxa. Interestingly, the 
tnT-trnF region was estimated to be 1800 bp, 1850 bp 
and 2000 bp in length for S. nivalis, S. tenuis and S. 
foliolosa, respectively. 

Sixteen out of 105 enzyme restriction assays showed 
restriction site differences between S. foliolosa and 
the S. nivalis complex, including three site differences 
in the trnH-trnK region, three in the rbcL-ORF106 
region, three in the trnV-rbcL region, five in the trnF- 
trnV region and two in the trnT-trnF region (Table 
4). No restriction site differences were detected bet- 
ween S. nivalis and S. tenuis. However, digestion of 
the trnT-trnF region with the restriction enzyme Alul 
showed clearly the length difference between S. 
nivalis and S. tenuis (Figure 2). 


Table 4. Data matrix of restriction site characters for S. nivalis (N), S. tenuis (T) and S. foliolosa (F). “0” represents absence 


of restriction site; “1” represents presence of restriction site. 


trnH-trnK rbcL-ORF106 
NTF INGE 
Alu I eal ie 1b kl 
Apal 000 000 
BamHI 000 eeleall 
Bell 000 000 
Bel Il 000 Ave let 
BstO I 001 Ethel 
Dral 001 lO 
EcoRI 000 Ikea 
EcoR V 000 ae al 
Hae It Hee al 001 
Hha lI 000 000 
Hinf I lbw eal 
Msp I ha al hiked 
j sO lal al 
apna iL hal 
000 000 
000 Write al 
Pal il ilies al 
ie ihel oak il 
000 001 
000 000 


trnT-trnF 
NTF 


trnF-trnV 
INiglge 


trnV-rbcL 
NTE 


DR RP eB RrP ORR RRP rR OOOO COOCCrF 
ORB Re RP ROR RFP RRP rR OCOCOCOCOCCOOF 
CORR RRB BS OR RP RB Re RO OrR KR OOF OF 
eee Re RK Or Fer COOCOCcoocoocoorcoor 
[St ee a) ees) (5) oa ea) (Fea) fos 
OPER ROORHEH OHM OH COOCOOH 


SPSS LOLS shore Oise SO oro Sc 
See =O Ores Orr Ore SO Ora Ore SS 
OororcjooroqooqococrreKScocorf 


Oo 
Oo 
Nw 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


MNNTTTTNNT F 


FiGuRE 2. Digestion of the chloroplast DNA region trnT-trnF with restriction enzyme Alul for Saxifraga nivalis (N), S. 
tenuis (T) and S. foliolosa (F). The first lane contains a 100 bp DNA size marker (M). Note absence of the upper band 


on S. nivalis. 


Discussion 

The principal component analysis confirms that the 
S. nivalis complex in the Canadian Arctic is represent- 
ed by two principal morphotypes, appearing as adjacent 
clusters on the PCA graph. The several plants occu- 
pying an intermediate position between the two clus- 
ters displayed a somewhat intermediate morphology, 
but nevertheless could readily be assigned to one of 
the two taxa. For example, several specimens of S. 
nivalis from Prince Patrick Island (e.g., Gillespie & 
Consaul 6869-4, 6926a-3) were very small with slender 
flowering stems, characters typical of S. tenuis, but 
otherwise had the morphological characteristics of S. 
nivalis. Although character states or ranges overlapped 
for the majority of characters examined, differences 
between the two taxa were statistically significant for 
all quantitative and for three qualitative characters. 


Morphological variability 

A significant amount of variance was observed for 
most characters measured. Since the S. nivalis complex 
is considered an environmental indicator (Aiken et 
al. 1998, 2000*), the variability observed may be due 
to phenotypic plasticity and variation in environment. 
For instance, S. nivalis plants growing in harsher 
habitats are usually shorter and less robust (e.g., the 
very short plant shown in Figure 3) than those grow- 


ing in more nourishing environments. Also plants of 
S. nivalis may be taller with more slender stems in 
shady microhabitats as a result of etiolation. Another 
contributor to variance is the time of collection. Since 
stems elongate in the fruiting stage, particularly in S. 
nivalis, plants collected later in the season are more 
likely to be taller than their younger counterparts. 
However, most examined collections of S. nivalis and 
S. tenuis made at the same time and site displayed 
significant height differences on average between the 
two taxa. Leaf characters, such as leaf size and teeth 
number and size, showed much variation both within 
and between the two taxa. Although S. tenuis is usu- 
ally found in wetter habitats than S. nivalis, the latter 
species in particular occupies a broad range of habitats 
and the two species sometimes co-occur in mixed 
populations. Some of the morphological variation in 
the complex, particularly within S. nivalis, may reflect 
this variation in habitat (Blondeau and Cayouette 2002; 
Aiken et al. 2000*). Such high intraspecific variation 
is apparently typical of Saxifrages, particularly in arctic 
and alpine environments, and several studies have 
demonstrated that such variation may not be of taxo- 
nomic significance (Soltis et al. 1996; Brysting et al. 
1996). 

The coloration of petals also varied within each 
taxon, but season and environment can be a large con- 


HEALY AND GILLESPIE: ALPINE SAXIFRAGE COMPLEX 


pws 


ki, 
y 
~ 


FIGURE 3. Saxifraga nivalis complex in the Canadian Arctic. A. S. tenuis (Gillespie & Consaul 6978-2), habit. B. S. nivalis 


(Gillespie and Consaul 6977-1), habit. C. Comparison of basal rosette of leaves of S. tenuis (on left, Gillespie & 
Consaul 6978-2) and S. nivalis (on right, Gillespie and Consaul 6977-1). A and B are the same scale. 


334 


tributor to petal coloration. Plants growing in more 
open, sunnier environments tend to have more pink 
or dark red anthocyanin pigment throughout the plant, 
including the petals. White petals may also sometimes 
turn pinkish with age. In contrast, little variation was 
observed within each taxon for the texture and color- 
ation of stem hair, and these characters were signifi- 
cantly different between taxa. 

Among traits traditionally used for identification of 
the S. nivalis complex, many displayed considerable 
intrataxon variance, with ranges often overlapping be- 
tween taxa. Our results also indicate a greater degree 
of variability within S. nivalis than in S. tenuis in the 
Canadian Arctic. This pattern of variation results in a 
somewhat obscure morphological boundary between 
the two species and has led to difficulties in construct- 
ing good identification keys. 


Characters and taxonomic identification 

Despite the considerable intraspecific variation, 
most commonly used identification characters were 
significantly different between the two taxa. Indeed, 
plants of S. nivalis were overall larger and more robust 
than S. tenuis, and the latter often had a more reddish 
tinge on the petals and stem hair, as previously found 
by numerous authors (Simmons 1906; Polunin 1940; 
Porsild 1957; Boécher et al. 1968; Léve 1983; Blon- 
deau 1989a; Webb 1993). 

Flowering stem width was an excellent identifica- 
tion character apart from a few individuals, with S. 
tenuis characterized by a width of 1 mm or less, con- 
sistent with Webb’s (1993) use of it as a key character. 
Consistently used by various botanists (e.g., Webb 
1993), stem hair characters were among the most use- 
ful for identification. Saxifraga nivalis usually has a 
much more conspicuous, longer, denser, coarse crisped 
white vestiture, while S. tenuis has a less conspicuous, 
shorter, finer, usually glandular pubescence. Canadian 
Arctic S. tenuis was found to always have stem hairs, 
thus not conforming to descriptions of the taxon in 
Greenland and Svalbard as having mostly glabrous 
stems (Bécher et al. 1968, 1978; Ro@nning 1996). 
Generally, S. nivalis plants were taller than S. tenuis, 
but due to its variance, height was not useful for iden- 
tification. 

Saxifraga nivalis had more numerous teeth on the 
leaves than S. tenuis, and the latter had rounder ob- 
tuse teeth versus more pointed, as suggested in the 
literature (e.g., B6cher et al. 1968), The width and 
length of S. tenuis teeth were also found to be less 
than those observed on S. nivalis, but, this is not sur- 
prising considering that S. nivalis had on average 
longer and wider leaves. Interestingly, S$. nivalis had 
a notably less regular dentition than S. tenuis in regards 
to the size of the teeth. The former generally had 
small teeth at the base with teeth becoming larger 
towards the apex, whereas S. tenuis had more evenly 
sized teeth. Although size of the largest leaf differed 
significantly between the two taxa, leaf size characters 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


make poor diagnostic characters due to overlap and 
age related variation. Also, leaf shape varied tremen- 
dously and thus should not be considered a good 
discriminating character. However, the leaf apices of 
S. tenuis did tend to be more rounded than those of S. 
nivalis. 

Inflorescence compactness provides a useful identi- 
fication character, with S. nivalis having one or more 
compact head-like clusters and S. tenuis a more open 
inflorescence (as described in Porsild 1957). Flower 
pedicels, especially of the lowermost flowers, are longer 
in S. tenuis than in S. nivalis. Since pedicels of the 
latter may elongate somewhat in fruit, this difference 
may be obscured later in the season. Although not con- 
stant, petals tend to be whiter, wider and ovate in S. 
nivalis and pink, narrower, more elongated and rounder 
at the apex in S. tenuis. BOcher (1938), Bocher et al. 
(1968), and more recently Love (1983) and Rgnning 
(1996) state that S. nivalis has straight or slightly 
spreading styles, while S. tenuis has styles that curve 
strongly downwards. This was not always the case in 
Canadian Arctic specimens we examined. The way 
specimens were pressed and the time of collection 
might be partly responsible for the variation in this 
character within the two species. 

Other characters showed variation that was not use- 
ful in separating the two taxa. While S. nivalis usually 
had hairs on leaf margins, this character was also 
found to a lesser degree in S. tenuis. Most specimens 
examined had a purplish or reddish stem, thus not 
specific to S. tenuis. Also, the flowering stems of both 
taxa had on occasion a reduced leaf (bract) or two near 
the inflorescence, although usually more prominent 
in S. tenuis. 

The presence of rust-coloured hairs on the lower 
surface of the leaves of S. tenuis has been commonly 
used to distinguish it from S. nivalis in the Canadian 
Arctic, and was used as a key character by several 
authors (Porsild 1957; Scoggan 1978; Porsild and 
Cody 1980). However, this study reveals that hairs 
are present on the leaf undersurface in over 99% of the 
plants examined of both species in the Canadian Arctic, 
and that hairs may be rust-coloured in both species, 
about 42% of S. nivalis plants examined and 74% of 
S. tenuis plants. This error may have resulted from 
Porsild’s inclusion of S. rufopilosa within his concept 
of S. tenuis (distribution maps of S. tenuis in Porsild 
(1957) and Porsild and Cody (1980) appear to have 
included S. rufopilosa). Saxifraga rufopilosa, a species 
now considered to be endemic to unglaciated areas of 
the Yukon and Alaska, is characterized by a mat of long 
crinkly rusty hairs on the leaf undersurface (Cody 
1996), much longer and denser than in either S. tenuis 
or S. nivalis. Whatever its origin, use of rust-coloured 
hair presence as a key diagnostic character in identify- 
ing Canadian Arctic material of the S. nivalis complex 
resulted in many misidentified specimens and much 
confusion as to the identity of S. tenuis. 


2004. HEALY AND GILLESPIE: ALPINE SAXIFRAGE COMPLEX 335 


Sy 


e Specimen data (CAN, DAO) 


‘\ == Adjacent species range from 
S Porsild and Cody (1980) 
we 


e Specimen data (CAN, DAO) 


-\se=, Adjacent species range from 
OB Porsild and Cody (1980) 


Ficure 4. Distribution of the Saxifraga nivalis complex in the Canadian Arctic. (above) S. nivalis. (below) S. tenuis. 
Shading indicates approximate distribution outside of the Arctic Islands (from Porsild and Cody 1980, and CAN and 
DAO herbarium records). 


336 


The many herbarium collections having both S. 
nivalis and S. tenuis plants on the same sheet and under 
the same number (e.g., Bruggeman 358, Savile 4562, 
Macdonald 150, Calder et al. 24149) shows just how 
problematic identification of this complex in the Cana- 
dian Arctic has been. About 20% of herbarium speci- 
mens identified as S. tenuis at both CAN and DAO 
were in fact mixed collections including both S. tenuis 
and S. nivalis plants, or less often including small plants 
of S. foliolosa or S. hieracifolia Waldst. & Kit. Pre- 
vious identification problems of the S. nivalis complex 
in the Canadian Arctic may also have been due to 
misinterpretation of key characters. For example, S. 
nivalis with several flower heads could be keyed out 
under lax (open) inflorescence. 

Although S. nivalis and S. tenuis can be readily dis- 
tinguished in the Canadian Arctic, a small percentage 
of specimens had one or a few characteristics of the 
other species. For example, some high arctic S. nivalis 
had very slender stems or fruit with strongly recurved 
styles. While this may be part of the natural variation 
within S. nivalis, it may also suggest a low level of 
hybridization. Some of the overlap in quantitative char- 
acter states shown in our morphological analyses may 
also be the result of hybridization. Recent cytological 
studies have suggested the existence of S. nivalis-S. 
tenuis hybrids (2n = 40) in regions of the Russian 
Arctic and the Yukon where S. nivalis and S. tenuis are 
found growing together (Krause and Beamish 1973; 
Devyatov et al. 1997). The population in the Gaspé 
Peninsula has the same intermediate chromosome num- 
ber and may represent a stabilized relictual hybrid in 
an area whether neither parent now occurs (Gervais et 
al. 1995). In addition Polunin (1959) suggested that 
hybridization may also occur between the S. nivalis 
complex and other closely related species, such as S. 
hieracifolia. 


Geographical variation in Saxifraga tenuis 

Savile (1961) proposed that North American S. tenuis 
may be divided into three disjunct and morphologi- 
cally distinct “populations.” The first is a high arctic 
population composed of plants identical to material 
from the north of Greenland and hypothesized to have 
spread from there following the Pleistocene. The 
second population is apparently found mostly along 
Hudson Straight in southern Baffin Island, northern- 
most Quebec and Southampton Island, but also 
scattered northward to northeast Baffin Island. Savile 
treated plants in the Gaspé Peninsula of Quebec, con- 
sidered by some as a separate species S. gaspensis, as 
part of this population. Plants of this population appar- 
ently tend to be larger and less purplish in colour than 
in the high arctic. The third highly variable population, 
found in Alaska and the Yukon, is usually character- 
ized by a highly purplish pigmentation and whitish 
scape hairs, and was suggested to be intermediate 
between S. nivalis and high arctic S. tenuis. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Our analysis indicates that eastern and high arctic 
plants of S. tenuis cannot be distinguished based on 
examined quantitative characters, while western mon- 
tane plants (based on our small sample size) may be 
differentiated from most arctic plants based on larger 
size. Since Savile (1961) used mostly qualitative char- 
acters to distinguish the three groups, further analyses 
based on both qualitative and quantitative characters 
and including more western montane and eastern arctic 
specimens should be made to further test his hypoth- 
esis. 


Chloroplast DNA analysis 

The five non-coding cpDNA regions of S. nivalis 
and S. tenuis were identical for restriction sites as- 
sayed. Considering the conservative nature of cpDNA 
evolution, it is not surprising to observe such similar- 
ity, with closely related species often having identical 
cpDNA restriction site profiles (Olmstead and Palmer 
1994). However, S. nivalis and S. tenuis displayed an 
estimated 50 bp length difference in the trnT-trnF 
region. This difference was observed both among plants 
from the same site and from different locations. This 
suggests that the size variation in the t7T-trnF region 
is not background variation among populations or a 
phenomenon limited to a particular population, but 
represents a molecular difference between S. nivalis 
and S. tenuis, either a deletion event in S. nivalis or 
an insertion event in S. tenuis. Sequencing of the 
tnT-trnF region would prove useful in specifying the 
nature of this size difference. The RFLP analysis re- 
vealed molecular differences for 16 of 105 restriction 
enzyme assays between S. foliolosa and the other two 
species. These results are similar to a study of Saxifraga 
based on the cpDNA matK sequences (Soltis et al. 
1996; plus sequences in Genbank), in which S. nivalis 
and S. tenuis would appear to be more closely related 
to each other than to S. foliolosa. Based on comparison 
of the Genbank sequences, the single plant of S. fenuis 
differed from S. nivalis in four nucleotide substitu- 
tions, and from S. foliolosa in 79 substitutions. 


Implications 

Our results suggest that S. nivalis and S. tenuis 
should be recognized as distinct taxa, and are most 
appropriately recognized at the species level following 
Porsild (1957), Porsild and Cody (1980), and Webb 
(1993). The two taxa are distinguishable at both the 
molecular and cytological level and have morpho- 
logical differences that correspond to these differences. 
Although no morphological characters give 100% sep- 
aration, many quantitative and qualitative characters 
show statistically significant differences between the 
two taxa. The two taxa overlap in geographic distribu- 
tion in the Canadian Arctic, with the range of the less 
common S. tenuis included within that of S. nivalis. 
Although ranges of habitat preferences differ, there is 
considerable overlap, with the two taxa sometimes oc- 
curring in mixed populations. Lack of geographical 


2004 


separation and considerable habitat overlap make 
recognition at the subspecific level inappropriate, while 
the morphological distinctiveness of the two taxa would 
make recognition at the varietal level inappropriate. 


Saxifraga nivalis complex in the Canadian Arctic: 
key to species 

A. Inflorescence one to several dense head-like clusters 

of numerous flowers; petals white or essentially so 

(sometimes becoming pink with age); flowering stem 

(0.5-) 1-2.5 mm wide, moderately to densely hairy, 

with conspicuous long coarse white hairs; plant usu- 

ally robust in appearance S. nivalis 

B. Inflorescence an open cyme of fewer flowers, flowers 

on distinct pedicels; petals pink or less often white; 

flowering stem 0.3-1 mm wide, usually sparsely hairy, 

with short fine hairs that are usually inconspicuous; 

plant delicate in appearance S. tenuis 


Saxifraga nivalis L. 

Plants perennial herbs, (2) 5—17 cm high, with a basal ros- 
ette of leaves. Leaves simple, alternate, all basal in a rosette, 
evergreen; petiole usually distinct, 0.2-2.5 mm long. Leaf 
blade ovate, obovate, orbicular or spatulate, 6-36 mm long 
(mean 14 mm), 4-21 mm wide (mean 10 mm), slightly leath- 
ery; green above, green or reddish-purple beneath, base obtuse 
and usually abruptly narrowed at petiole; apex obtuse, rounded 
or rarely acute; appearing single-veined or with inconspic- 
uous lateral veins; upper surface glabrous; lower surface 
sparsely to densely hairy; hairs 0.2-1.2 mm long, rust-coloured, 
whitish or translucent; margins coarsely serrate or crenate, 
usually with rust-coloured or whitish hairs; teeth 7-14, usually 
increasing in size towards blade apex, tips usually pointed. 
Flowering stem erect, (0.5) 1-2.5 mm wide, green to purple, 
moderately to densely hairy, with hairs conspicuous, (0.5) 
0.7-1.2 mm long, crisped (crinkly and irregularly wavy), white 
or translucent with tips and crosswalls white or sometimes 
rust-coloured or purplish, the tips sometimes glandular, usually 
leafless or occasionally with reduced leaves (bracts) close to 
the inflorescence. Inflorescence a dense head-like cluster of 
numerous flowers, sometimes with 1-2 smaller lateral clusters 
on short to long peduncles; flowers on very short pedicels, 
subtended by bracts. Flowers: sepals 5, free, green or purple, 
glabrous; petals 5, ovate or elliptic, 0.7-1.8 mm wide, equal 
to or longer than the sepals, white or cream-coloured, some- 
times with reddish apex, mostly obtuse at apex; stamens 10; 
gynoecium partly inferior; carpels 2, partly fused; styles 2, 
free; placentation axile; ovules numerous. Fruit a capsule, 
spherical in lower half, with straight or slightly divergent 
free carpels above, 34 mm long, 4.5—5.5 mm wide, glabrous, 
dry, dehiscent; styles straight or slightly divergent, persistent; 
sepals persistent in fruit. Seeds numerous, 0.5—1 mm long, 
yellowish brown, with surface verrucose. Figure 3b, c; map 
4a. 


Saxifraga tenuis (Wahlenb.) Harry Sm. 

Plants perennial herbs, 2-9 cm high, with a basal rosette 
of leaves. Leaves simple, alternate, all basal in a rosette, ever- 
green; petiole usually indistinct, ~0.5—2.5 mm long. Leaf blade 
narrowly ovate, obovate or spatulate, 4-15 mm long (mean 
7 mm), 2-8 mm wide (mean 5 mm); slightly leathery, green 
to reddish-purple above, reddish-purple beneath; base obtuse 
or broadly attenuate and decurrent onto petiole; apex obtuse, 
rounded or rarely acute; appearing single-veined or with 
inconspicuous lateral veins; upper surface glabrous; lower 


HEALY AND GILLESPIE: ALPINE SAXIFRAGE COMPLEX 


337 
surface usually sparsely hairy, with hairs 0.1-0.4 mm long, 
rust-coloured, whitish or translucent; margins coarsely serrate 
or crenate, with sparse rust-coloured or whitish hairs or some- 


times glabrous; teeth 6-9, usually evenly sized, tips usually 
rounded. Flowering stem erect, 0.3-1 mm wide, usually dark 
purple, sparsely hairy, with hairs mostly inconspicuous, 0.1- 
0.3 (0.5) mm long, fine, crinkled or straight, rust-coloured 
to translucent, often with purplish crosswalls and tips, the 
tips usually glandular; sometimes with a reduced leaf (bract) 
on the upper part of the stem. Inflorescence an open head of 
few flowers; flowers subtended by bracts; lower-most flowers 
often long-pedicellate and each subtended by a conspicuous 
bract. Flowers: sepals 5, free, green or purple, glabrous; petals 
5, ovate or elliptic, 0.5-1.1 mm wide, equal to or longer than 
the sepals, usually pink, sometimes white, often with reddish 
apex, often rounded at apex; stamens 10; gynoecium partly 
inferior; carpels 2, partly fused; styles 2, free; placentation 
axile; ovules numerous. Fruit a capsule, spherical in lower half, 
with diverging free carpels above, 3-4 mm long, 4.5—5.5 mm 
wide, glabrous, dry, dehiscent; styles usually strongly re-curved 
at maturity, persistent; sepals persistent in fruit. Seeds num- 
erous, 0.5—1 mm long, yellowish brown, with surface verru- 
cose. Figure 3a, c; map 4b. 


Acknowledgments 

We thank Kristina Makkay for assistance with the 
Statistical analysis and figures; Laurie Consaul for sug- 
gestions in statistical analysis and assistance with map- 
ping; Michelle Leblanc for assistance with mapping: 
Susan Aiken for interesting discussions; Gisele Mitrow 
and William J. Cody for assistance at the DAO herb- 
arium. Also, a special thank-you to Marcel Blondeau 
from the Université Laval for sharing his knowledge 
and insights on the Saxifraga nivalis complex. We 
acknowledge and thank Polar Continental Shelf Project, 
Nunavut Research Institute and Aurora Research 
Institute for their logistic support in the field. This 
paper is based on a fourth year honours thesis project 
by C. H. submitted to the Biology Department, Uni- 
versity of Ottawa. 


Documents Cited (marked * in text). 

Aiken, S. G., M. J. Dallwitz, L. L. Consaul, C. L McJannet, 
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2000. Flora of the Canadian Arctic Archipelago: Descrip- 
tions, Illustrations, Identification, and Information Re- 
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Aiken, S. G., M. J. Dallwitz, L. L. Consaul, C. L. McJannet, 
L. J. Gillespie, R. L. Boles, G. W. Argus, J. M. Gillett, 
P. J. Scott, R. Elven, M. C. LeBlanc, and A. E. Zamluk. 
2002. Flora of the Canadian Arctic Archipelago: Descrip- 
tions, Illustrations, Identification, and Information Re- 
trieval. (version 14 August 2002) http://www.mun.ca/ 
biology/delta/arcticf/ 


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Received 4 February 2003 
Accepted 7 December 2004 


Appendix 1: Collections examined of the Saxifraga nivalis complex from the Canadian Arctic. 


Saxifraga nivalis 

NUNAVUT. Axel Heiberg Island: 79°28'N 87°40'W, Beschel 10979 (CAN); Bukken River, 80°31.57'N 92°21.83'W, 
Gillespie et al. 6613 (CAN); Crusoe Glacier, 97°27'N 91°12'W, Kuc 476-3 (CAN); Crusoe River, 79°24'N 90°S0'W, Kuc 
s.n. (CAN 331187); Diana Lake, 79°30°N 88°30’ W, Porsild 16699-2 (CAN); Expedition Fjord, 79°20'N 92°W, Kuc s.n. 
(CAN 331189, CAN 331189); Mesa Brook, 79°37'N 95°W, Beschel 13129 (CAN); Mokka Fiord, 79°45'N 87°W, Parker 
73061E (CAN). Baffin Island: 63°24'N 64°35'W, McLaren 91 (CAN); 66°40'N 70°W, Soper s.n. (CAN 65150); Admiralty 
Inlet, 73°13'N 84°W, Malte 118993 (CAN); Apex, 63°43'N 68°27'W, Gillespie & Soreng 6785 (CAN); Beekman Peninsula, 
63°24'N 64°35'W, McLaren 9] (CAN); Burwash Bay, 65°59'N 71°18'W, Jacobs & Maus s.n. (CAN 517697); Frobisher 
Bay, 63°45'N 68°31'W, Gillett 19018 (CAN); Dorset Island, 64°12'N 76°32'W, Hainaud & Norman 5698 (CAN); Foxe 
Peninsula, 66°23'N 83°13'W, Manning 265 (CAN); Foxe Peninsula, 69°52'N 76°54'W, Manning 231 (CAN); Frobisher Bay, 
62°57'N 66°03'W, Aiken et al. 86-260 (CAN); Frobisher Bay, 63°44'N 68°27'W, Aiken et al. 86-493 (CAN); Frobisher Bay, 
62°57'N 66°03'W, Calder et al. s.n. (DAO 24148); Frobisher Bay, 62°57'N 66°03'W, Wynne-Edwards 7353 (CAN); Pond 
Inlet, 72°50'N 76°40'W, Dutilly 1224a (CAN); Stising Valley, Weber 94 (CAN); Taverner Bay, 67°12'N 72°25'W, Manning 
44, 68, 77 (CAN). Bathurst Island: Polar Bear Pass, 75°43'N 98°23'W, Aiken & Maus 92-022 (CAN); Goodsir Inlet, 
75°45'N 97°45'W, Lamothe 69-25 (CAN); Polar Bear Pass, 75°43.8'N 98°25.45'W, Gillespie & Consaul 6984 (CAN) [DNA 
voucher, individuals 1-5 examined]. Bylot Island: 72°53'N 76°W, Wilcox 125631 (CAN). Cameron Island: 76°19.5'N 
104.5°W, Aiken 92-039 (CAN). Cornwallis Island: Resolute, 74°40'N 94°50'W, Aiken & MacCormac 98-055 (CAN 
581909, plants 1-5; CAN 582554); Resolute, 74°40'N 95°04'W, Aiken et al. 93-072 (CAN). Devon Island: Dundas 
Harbour, 74°31.3'N 82°33.5'W, Gillespie et al. 6675-1 (CAN); Truelove Lowland, 75°40'N 84°40'W Kerik s.n. (CAN 
409363). Digges Island: 62°32'N 77°45'W, Gaston 25, 29 (CAN); 62°35'N 77°50'W, Gaston 17 (CAN). Ellef Ringnes 
Island: Isachsen, 78°47'N 103°50'W, Macdonald 242 (CAN). Ellesmere Island: 79°34'N 84°44'W, Edlund & Roncato- 
Spencer 264 (CAN); Alexandra Fiord, 78°53'N 75°45'W, Gillespie & Vogel 6165 (CAN); Alexandra Fiord, 78°53'N 
75°50'W, Gillett & Shchepanek 18108 (CAN); Craig Harbour, 76°20'N 81°30'W, Dutilly 1267 (CAN): Hazen, 68°30'N 
72°45'W, Soper 8194 (CAN); Hazen, 81°49'N 71°20'W, Gillespie & Vogel 6238 (CAN); Sawtooth Range, 79°43.54'N 
83°09.44'W, Gillespie et al. 6643 (CAN); Slidre Fjord, 80°N 86°15'W, Tener 95 (CAN): Tanquary, 78°53'N 75°45'W, 
Gillespie & Vogel 5982 (CAN). King William Island: Gjoa Haven, Cooper 28 (CAN), 73 (CAN), 226 (CAN); Gjoa 
Haven, 68°38'N 95°53'W, Larson 25 (CAN); Victory Point, Cooper 164 (CAN), 122 (CAN). Lougheed Island: 77°30'N 
105°38'W, Edlund 2091 (CAN). Meighen Island: 79°55'N 99°30'W, Kuc (CAN 331171). Pelly Bay: 68°53'N 89°51'W, 
Campbell s.n. (CAN 282778). Prince Charles Island: 67°10'N 76°43'W, Baldwin 1919 (CAN). Victoria Island: 68°53'N 
105°W, Edlund & Argus 12730-1 (CAN). 

NORTHWEST TERRITORIES. Banks Island: 73°N 117°W, Aiken 99-203 (CAN); 73°24'N 117°0'W, Porsild 17711 
(CAN); 74°31'N 121°07'W, Manning & Macpherson 164 (CAN); Bernard River, 73°22'N 121°47'W, Maher & Maclean 75 
(CAN); Egg River, 72°27'N 124°36'W, Lambert s.n. (CAN 535943); Lambton, 71°05'N 123°09'W, Porsild 17579 (CAN); 
Shoran Lake, 71°51'N 113°23'W, MaclInnes s.n. (CAN 535570, 535598, 535733). Fitzwilliam Owen Island: 77°07'N 
113°47'W, Kuc s.n. (CAN 331114). Melville Island: 75°56'N 114°48'W, Edlund & Aiken 148 (CAN); Canrobert Hills 
75°47.30'N 115°56'W, Edlund 179 (CAN); Ibbett Bay, 75°54'N 114°30'W, Edlund 43 (CAN); Kitson River, 76°02'N 
113°05'W, Edlund 18 (CAN); Marie Bay 76°12'N 114°55'W, Edlund 305 (CAN); McCormick Inlet, 75°46.2'N 
112°31.64'W, Gillespie & Consaul 6977 (CAN, plants 1, 2), 6964a (CAN); Shellabear Point, 71°51'N 113°23'W, Edlund 
115 (CAN); Sherard Bay, 76°08'N 108°07'W, Dugal s.n. (CAN 535598); Winter Harbour, 74°17'N 110°42'W, Kuc s.n. 
(CAN 400104); Winter Harbour, 74°17'N 110°42'W, Tener & Harington 149 (CAN). Prince Patrick Island: Green Bay. 
76°33.67'N 118°53.09'W, Gillespie & Consaul 6869 (CAN, plants 1-6); Intrepid Inlet, 76°22.01'N 118°35.45'W, Gillespie 


340 THE CANADIAN FIELD-NATURALIST Vol. 118 


& Consaul 6876 (CAN); Mould Bay 76°12N 119°25'W, MacDonald 146 (CAN); Mould Bay, 76°14.50'N 119°21'W, 
Gillespie & Consaul 6806 (CAN, plants 1-5); Mould Bay, 76°14'N 118°57'W, Gillespie & Consaul 6827 (CAN, plants 1, 
2), 6826 (CAN, plants al-a5); Mould Bay, 76°21.50'N 119°30.50'W, Gillespie & Consaul 6884 (CAN, plants 1, 2), 6984 
(CAN, plants 1-5), 6806 (CAN, plants 1-5). Victoria Island: Minto, 71°34'N 115°21'W, Edlund 594 (CAN). 


Saxifraga tenuis 

NUNAVUT. Axel Heiberg Island: Bastion Ridge, 79°21'N 90°48'W, Beschel 12920 (CAN); Mesa Brook, 79°35'N 
95°30'W, Beschel 13129 (CAN 295548): 80°17'N 88°27'W, Scotter & Zoltai 45133 (DAO), 4518/1 (DAO). Baffin Island: 
Resolution Island, 61°20'N 64°55'W, Wynne-Edwards 7225 (CAN); Erik Harbour, 72°40'N 76°30'W, Calder s.n. (DAO 
30869, 30871); Ogac Lake, 62°51.7'N 67°21'W, McLaren s.n. (CAN 274125); Frobisher Bay, 63°45'N, 68°32'W, Senn 3658 
(DAO); Blackhead Island, 64°59'N 66°19'W, Soper s.n. (CAN 125598); Savage Islands, 61°49.15'N 65°42.62'W, Gillespie 
et al. 6728 (CAN), 6732 (CAN); Inugsuin Fjord, 69°50'N 69°08'W, Beschel 3649A (CAN). Bathurst Island: 76°36'N 
100°04'W, Blake 24a (DAO). Bylot Island: 73.03'N 80.07'W, Scotter & Zoltai 6751A (DAO). Ellef Ringnes Island: 
Isachsen, 78°49'N 103°37'W, Savile 4275 (DAO), 4219 (DAO), 4224 (DAO), 4285 (DAO), 4355 (DAO), 4293 (DAO); 
Isachsen, 78°47'N 103°30'W, MacDonald 251 (CAN), 252 (CAN); Christopher Peninsula, 78°59'N 101°35'W, Savile 4194 
(DAO); 78°45'N 102°50'W, Savile 4148 (DAO). Ellesmere Island: Eureka, 79°59'N 85°50'W, Scotter & Zoltai 45294-B 
(DAO); Eureka, 80°10'N, 85°39'W, Bruggemann S05 (DAO); Fosheim Peninsula, 79°34'N 84°44'W, Edlund & Roncato- 
Spencer 262 (CAN 535348, 533280); Fosheim Peninsula, 79°58'N 84°26'W, Edlund & Roncato-Spencer 275 (CAN); 
Hazen, 81°49'N 71°21'W, Savile 4485 (DAO), 4752 (DAO), 4655 (DAO); Hazen, 81°45'N 68°30'W, Powell 338 (CAN); 
Hazen, 82°N 70°W, Soper 8273 (CAN); Makinsen Inlet, 76°41'N 81°37'W, Blake 7 (DAO); Mount Pullen, 82°25'N 
62°17'W, MacDonald 46 (CAN); 81°45'N 62°11'W, Powell 520 (CAN); Mt. Pullen, 82°26'N 62°11'W, Bruggemann 215 
(DAO); Tanquary Fjord, 79°12'N 83°29'W, Brassard 3306 (CAN); Van Hauen Pass, 81°07'N 86°55'W, Brassard 3047 
(CAN); Vesle Fjord, 79°12'N 83°29'W, Edlund & Roncato-Spencer 282 (CAN). Somerset Island: 72°56'N 94°57'W, Zoltai 
741044 (DAO). 

NORTHWEST TERRITORIES. Melville Island: 74°58'N 115°02'W, Parker 44f (DAO); Bailey Point, 74°58'N,115°02'W, 
Parker 48b (DAO); McCormick Inlet, 75°46.2'N 112°31.64'W, Gillespie & Consaul 6978 (CAN, plants 1, 2), 6964b 
(CAN). Prince Patrick Island: Green Bay, 76°33.67'N 118°53.09'W, Gillespie & Consaul 6870 (CAN, plant 1); Intrepid 
Inlet, 75°46.9'N 112°22.09'W, Gillespie & Consaul 6877 (CAN, plant 1); Mould Bay, 76°21.5'N 119°30.5'W, Gillespie & 
Consaul 6881 (CAN, plants 1-6); Mould Bay, 76°14'N 118°57'W, Gillespie & Consaul 6825 (CAN, plants 1-3); Mould 
Bay, 76°14.5'N 119°21'W, Gillespie & Consaul 6807 (CAN, plants 1-5). Victoria Island: Ulusartok Peninsula, 70°44'N 
117°44'W, Edlund 875 (CAN). 

ALASKA. Kenai Peninsula. 60°49'N 149°33'W, Calder 6511, 6249, 6107 (DAO). 

BRITISH COLUMBIA. Haines Junction, Calder & Kukkonen 28151 (DAO); Haines, Taylor et al. 1204 (DAO). 


Mixed collections - Saxifraga tenuis and Saxifraga nivalis on same sheet 

NUNAVUT. Baffin Island: Apex, 63°43'N 68°27'W, Calder et al. s.n. (DAO 24149); Home Bay, 68°45'N 67°10'W, Smith 
VP-53-61 (CAN). Ellesmere Island: Tanquary Fjord, 79°12'N 83°29'W, Brassard 3317 (CAN); Hazen, 81°49'N 71°21'W, 
Savile 4562 (DAO). 

NORTHWEST TERRITORIES. Victoria Island: near Holman, 70°43'N 117°43'W, Edlund 719 (CAN). Prince Patrick 
Island: Mould Bay, 76°12'N 119°25'W, MacDonald 150 (CAN, S. tenuis: 1, 3, 4, 7, 8, 9; S. nivalis: 2, 5, 6, 10). 


_ The Influence of Prey Availability and Vegetation Characteristics on 
| Scent Station Visitation Rates of Coyotes, Canis latrans, in a 
Heterogeneous Environment 


LynpDA A. RANDA! and JOHN A. YUNGER2 


Department of Biological Sciences, Northern [Illinois University, DeKalb, Illinois 60115 USA 

'Present address: Natural and Applied Sciences Division, College of DuPage, 425 Fawell Boulevard, Glen Ellyn, Illinois 
60137 USA 

Present address: Environmental Biology Program, Governors State University, University Park, Illinois 60466 USA 


Randa, Lynda A., and John A. Yunger. 2004. The influence of prey availability and vegetation characteristics on scent station 
visitation rates of Coyotes, Canis latrans, in a heterogeneous environment. Canadian Field-Naturalist 118(3): 341-353. 


We investigated the effects of local prey fluctuations and habitat variables on the scent station visitation rates of the Coyote 
(Canis latrans) in northern Illinois within a heterogeneous environment. Availability of small mammalian prey was assessed 
by monthly mark-recapture sampling and visual counts conducted along three, 192-m transects in each of seven habitats that 
ranged from grassland to wooded sites. Habitat metrics, which included foliage density, ground cover, and canopy cover, 
were also collected for the same seven habitats. Visitation rates of Coyotes were determined from scent station lines parallel 
to the small mammal trapping transects. A multiple regression analysis indicated that Coyote visitation rates across the study 
site were influenced positively by vole (Microtus spp.) abundance and negatively by canopy cover. When Coyote visitation 
rates were regressed on vole abundance for only the habitats in which voles occurred, the relationship was not significant. 
This may be attributed to the general avoidance of wooded areas by Coyotes. Coyotes did, however, respond to experimentally- 
induced abundant patches of Peromyscus. These findings suggest Coyotes selectively use grassland habitats within a hetero- 


geneous environment and may modify their use according to prey availability. 


Key Words: Coyote, Canis latrans, habitat use, heterogeneous environment, prey availability, scent station, Illinois. 


The Coyote, Canis latrans, is among the most adapt- 
able of North American predators. Across an expan- 
sive range, the species occupies a variety of habitats 
and consumes a diversity of foods (Bekoff 1978). As 
such, the Coyote has been the focus of various local 
and regional studies of distribution, activity patterns, 
and diet selection. In particular, spatial and temporal 
activity patterns have been explicitly described (e.g., 
Andelt and Gipson 1979; Holzman et al. 1992), yet 
there has often been only circumstantial evidence as 
to causal factors (e.g., Person and Hirth 1991). Under- 
standing potential determinants of Coyote spatial dis- 
tribution and activity patterns requires an understanding 
of factors such as diet composition and specific use of 
available habitat types. 

Predators exhibiting flexible behavior patterns, such 
as Coyotes, provide an opportunity for studying possi- 
ble rapid responses to prey availability. Previous studies 
have shown that diets (Andelt et al. 1987: Brillhart 
and Kaufman 1995; Todd et al. 1981) and temporal 
activities of Coyotes (Shivik and Crabtree 1995) can 
vary seasonally based upon prey availability. Because 
space use and movements of vertebrate predators are 
closely coupled to foraging behavior, Coyotes could 
also alter their spatial use patterns in response to chang- 
es in prey availability. However, vertebrate predators 
frequently move over large spatial scales, whereas their 
smaller prey tend to be restricted to specific habitats 
or patches within a habitat. The study of prey in only 


one or a limited number of locations used by the pred- 
ators (Hamlin et al. 1984; Jaksic et al. 1993; Korsch- 
gen and Stuart 1972) may preclude accurate inferences 
on the foraging behavior of vertebrate predators. Thus, 
it is important to relate predator foraging behavior 
among habitats to prey abundances within habitats of 
heterogeneous landscapes (Dunk and Cooper 1994; 
Korpimaki 1994). 

In addition to prey, variations in vegetation structure, 
such as foliage density and height, among different 
habitats can also influence the movements and activities 
of vertebrate predators (Clark et al. 1993; Lamberson 
et al. 1994), especially in fragmented landscapes. 
Habitat differences may constrain predator foraging 
in certain areas through predators selectively using or 
avoiding particular habitat types (DeJong 1995; Small- 
wood 1995). As a result, predators may not respond 
to changes in prey availability within certain habitats. 
However, vegetation measures used to quantify pred- 
ator habitat use may also be related to or reflect prey 
usage (Anthony et al. 1981). Such inter-relatedness 
must be considered when making inferences of pred- 
ator behavior. 

We investigated the relationships of spatial and tem- 
poral distribution of prey and vegetation characteristics 
to the habitat visitation of Coyotes across a hetero- 
geneous environment. We incorporated experimental 
manipulations of habitat and prey, along with natural 
perturbations, to elicit predator behavioral responses 


341 


342 


within a relatively short time scale. Data collected on 
prey abundance and habitat variables were used to ad- 
dress whether scent station visitation rates of Coyotes 
vary among habitats and within a particular habitat in 
response to changes in prey availability. 


Study Areas 

The study site was located in northern Illinois at 
Fermi National Accelerator Laboratory (Fermilab) in 
Batavia, Illinois (41°50'N, 88°15'W). Fermilab encom- 
passes approximately 3200 ha and is characterized as 
a heterogeneous area, marked by distinct habitat bound- 
aries. The areas surrounding Fermilab consisted of 
small open spaces of mainly agricultural land inter- 
spersed with light industry and residential neighbor- 
hoods, providing potential predator access into or out 
of the study site. The study was conducted at seven 
different locations within Fermilab, representing five 
habitat types: (1) oldfield, dominated by Queen Anne’s 
Lace (Daucus carota) and Tall Goldenrod, (2) tallgrass 
prairie, dominated by Big Bluestem (Andropogon ger- 
ardii) and Indian Grass (Sorgastrum nutans); (3) brome 
grass field, dominated by Smooth Brome (Bromus 
inermis); (4) shrubby oldfield, a heterogeneous mix of 
dense shrubs and shrub-grassland mixture dominated 
by Tall Goldenrod (Solidago altissima) and Gray Dog- 
wood (Cornus racemosa) with small stands of Trem- 
bling Aspen (Populus tremuloides); and (5) oak wood- 
land, dominated by mature oaks (Quercus spp.), one 
woodland (Oak Woods I) characterized by less dense 
canopy (primarily Q. macrocarpa and Q. rubra) than 
the other (Oak Woods II; predominantly Q. alba and 
Q. rubra; Figure 1). These different habitats were cho- 
sen because they represented the heterogeneity of the 
study site and included all prey species potentially 
found in Coyote diets at Fermilab. A concurrent inves- 
tigation indicated that three prey, voles (primarily 
Meadow Voles, Microtus pennsylvanicus, and rarely 
Prairie Voles, M. ochrogaster), Eastern Cottontails 
(Sylvilagus floridanus), and mice (Peromyscus spp.), 
constituted 39.4%, 31.3%, and 8.4% of the Coyote 
diets, respectively, which was at least two times great- 
er than the proportion of any other prey item (Randa 
1996). 


Methods 
Availability of prey 

From March 1994 through May 1995, small mam- 
mal species compositions and abundances were esti- 
mated by mark-recapture live-trapping on three, 192 m 
transects located within each of the seven habitats 
(Figure 1). The transects were parallel and spaced from 
70 to 85 m to help maximize their independence in 
relation to small mammal movements. At least 50 m 
was maintained from each transect to habitat bound- 
aries to avoid edge effects. Seventeen trap stations were 
spaced at 12-m intervals along each transect, totaling 
51 trap stations per habitat. One 23 x 9 x 7.5 cm 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Sherman live-trap, baited with a peanut butter and 
oats mixture, was placed at each station. All habitats 
were trapped for two nights on a monthly basis. Traps 
were checked early morning and late afternoon (ca. 
every 12 hr). Captured individuals were marked with 
a uniquely numbered eartag (Monel Number 1, Na- 
tional Band and Tag Co., Newport, Kentucky). 

Nocturnal visual counts, using a high-powered spot- 
light (Woolf et al. 1993), were conducted for Eastern 
Cottontails in each of the seven habitats. The counts 
were conducted twice during each of the 15 small 
mammal trap sessions by sighting with binoculars, at 
one end and midway, along the small mammal trap- 
ping transects. Animals within ca. 25 m of the tran- 
sect were counted. For the visual counts, the trapping 
transects were lengthened to 250 m, delineated by flag- 
ging at opposing ends, to allow for the larger spatial 
movements of rabbits. Care was taken not to double 
count animals potentially moving between transects. 

For the experimental studies, fire was used as a 
means of manipulating prey. Tallgrass Prairie I was 
burned between the October and November 1994 
small mammal surveys, a period of time when small 
mammal abundances peak in northern Illinois (Yunger 
2002). This site was adjacent to Oak Woods I (Figure 
1), which concurrently experienced a high acorn 
production year. Both habitats contained two closely 
related species of small mammals (Peromyscus mani- 
culatus, the Deer Mouse, in the prairie and P. leu- 
copus, the White-footed Mouse, in the woods), which 
were similar in body size and morphology. Numerical 
responses, caused by immigration of Deer Mice to 
prairie burns, have been documented previously (Cook 
1959; Kaufman et al. 1983, 1988). Similarly, acorns 
have been shown to be an important local food source 
for mice (Peromyscus spp: Ostfeld et al. 1996; Wolff 
1996). Consequently, we predicted a localized increase 
of P. maniculatus in Tallgrass Prairie I and P. leucopus 
in Oakwoods I. The proximity of these two habitats 
provided the opportunity to compare changes in Coy- 
ote habitat visitation following the manipulations of 
prey abundance. 

Prey also was manipulated in Tallgrass Prairie II 
through food supplementation. From October 1994 
through March 1995, 10 kg of commercial rodent chow 
was hand broadcasted at weekly intervals on two of 
four (i.e., two treatment and two control), 0.60-ha 
square plots. As part of a separate study on the popu- 
lation dynamics of small mammals (Yunger 2002), 
prey numbers were determined by monthly trapping 
on a 6 X 6 grid with 12-m spacing in each plot. 
Predator spatial and temporal visitation responses to 
resulting changes in prey due to the food supplemen- 
tation experiment were also compared to the natural 
increase in acorns of Oak Woods I and to the prey 
response to experimental burning of Tallgrass Prairie I. 


Visitation rates of Coyotes 


2004 RANDA and YUNGER: SCENT STATION VISITATION RATES OF COYOTES 


es me mee 


n e le 
hu 


| 


= 


A 


( 


(|) Ibo! 


oO 


a 
senses 
“amas om sow 6 ame a 


: 1 | ear 
Bee oo 
Bea Pha 1 Oldfield 
Wea 2 Tallgrass Prairie I 
mane 3 Oak Woods I 


4 Shrubby Oldfield 
5 Oak Woods II 
6 Brome Grass Field 


10) 500 m Ace 
N ——— ee ransect 7 Tallgrass Prairie II 


FIGURE 1. Map of Fermilab showing the seven different study locations (habitats) and approximate locations of scent station 
transects in each habitat. Dashed line represents boundary of Fermilab. Roads, buildings, and open water (dark 


shaded areas) are included to further depict heterogeneity of the study site. 


344 


Coyote visitation rates were monitored through the 
use of scent stations. These were constructed by exca- 
vating a 50-cm diameter by 10-cm deep circular depres- 
sion, rimmed up to the ground surface with a 10-cm 
wide strip of aluminum flashing to inhibit invasive 
growth from surrounding vegetation, and filled with 
fine-grained sand. The resulting stations were level 
with the ground surface and fairly inconspicuous 
except for the lack of vegetation. Scent stations were 
located along 175-m transects, parallel to the small 
mammal trapping transects, in each of the seven loca- 
tions. Transects were spaced ca. 100 m apart while 
maintaining at least 50 m from a transect to the habi- 
tat edge. Eight scent stations were spaced every 25 m 
along each transect in the seven habitats, for a total 
of 24 scent stations per location. At the start of each 
monitoring period, all scent stations were baited by 
placing a cotton swab dipped in a liquid commercial 
predator lure (Cronk’s Predator 500, Wisscasset, Maine) 
upright in the middle of the scent station and the sand 
was smoothed. Stations were simultaneously monitored 
for a minimum of 2 nights (without precipitation), | 
to 3 times monthly (mean of 2.1 times per month); 
sand was not smoothed or otherwise disturbed by the 
authors during each monitoring period. Monthly scent 
station use was calculated for each habitat based upon 
visitation rates, or the proportion of scent stations 
exhibiting Coyote tracks in a given habitat, and divid- 
ed by the total number of operative nights to account 
for any variation in sampling duration. 

Within-habitat Coyote visitation rates were evaluated 
through the small mammal food supplementation 
experiment conducted in Tallgrass Prairie II. In Jan- 
uary 1995, one scent station was placed at each corner 
of the four plots previously described for this habitat. 
These 16 stations were monitored from February 1995 
through May 1995 to compare scent station visitation 
rates of Coyotes to prey numbers in supplemented 
and non-supplemented plots. 


Vegetation analysis 

Vertical foliage density, percent ground cover, and 
canopy cover were used to describe habitat structure 
for each of the seven habitats. Vegetation measurements 
were collected at 24 stations per habitat, along alter- 
nating stations of the small mammal trapping transects. 
Foliage density was measured using a modified profile 
board technique (MacArthur and MacArthur 1961). 
Percent cover of three, 30 x 50-cm profile boards, 
divided into 15, 10 x 10-cm cells, was measured at 
0.5 m, 1.0 m and 1.5 m from ground to mid-level of 
the board. A cell was considered covered if 50% or 
more of the cell, viewed from a 5-m distance, was ob- 
structed by vegetation. The proportion of cells covered 
determined percent cover of an individual board. All 
readings were taken facing north and there was no 
apparent patterning or trending in vegetation struc- 
ture and diversity among each of the four cardinal 
directions. To arrive at a single foliage density value 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


for each habitat, the proportion of vegetation cover 
was averaged across the three vertical profile boards 
for each sampling station, then the mean of the result- 
ing 24 values was calculated. 

Percent ground cover was assessed at each sam- 
pling station by the amount of vegetation (herbaceous, 
woody, and leaf litter) occupying a 1-m? quadrat against 
visible patches of ground (Brower et al. 1990). Canopy 
cover was estimated by the presence or absence (scored 
as | or 0, respectively) of vegetation viewed through 
a vertical ocular tube (James and Shugart 1970) at 
each sampling station. Average ground cover and 
canopy cover for each habitat was obtained by taking 
the mean of the 24 values of each metric. 

The vegetation analysis was conducted once in 
March 1994 and once in August 1994, representing 
vegetation structure characteristic of the non-growing 
and growing seasons, respectively. Two months of the 
year, May and November, represent transitions between 
the growing and non-growing seasons and hence, char- 
acteristic vegetation density. For this study, vegetation 
variables for May were described by the August sur- 
vey (i.e., high foliage density), and vegetation variables 
for November were described by the March survey 
(i.e., low foliage density). 


Data analysis 

Analyses of small mammal prey were based upon 
minimum number known alive (MNKA). This esti- 
mate was generated for each individual transect per 
month using Package C. M. R. (Le Boulengé 1987). 
The estimates then were averaged for the three tran- 
sects to yield monthly small mammal species compo- 
sition and abundance for each habitat. 

To examine whether Coyotes shifted visitation rates 
in response to prey, it was necessary to determine 
whether spatial and temporal differences in prey abun- 
dances existed. Analysis of covariance (ANCOVA), 
conducted with SAS PROC GLM (SAS Institute Inc. 
1990), was used to compare abundances of the most 
important prey species identified in Fermilab Coyote 
diets (Randa 1996). Specifically, analyses were con- 
ducted to test for significant fluctuations in prey abun- 
dances over time among each of the seven habitats. 
ANCOVA can be used to test for an interaction or 
heterogeneity of slopes (Littell et al. 1991) and was 
chosen because sampling units (i.e., habitats) were 
not replicated in space. Temporal changes of prey popu- 
lations were indicated by crossing the categorical vari- 
able (1.e., prey abundances across habitats or prey abun- 
dances per habitat) with the continuous covariate (time). 
Traditional model building techniques were used 
(Box et al. 1978) in which non-significant, higher- 
order interactions were removed from the model. If 
no significant sources of variation were detected, the 
full model was reported. If significant interaction of 
variables occurred (i.e., heterogeneity of slopes), sig- 
nificance of main effects were inferred from the plot- 
ted data. Statistical inferences were based on type III 


2004 


sum-of-squares and significance accepted at a = 0.05. 

Multiple regression (SAS PROC REG; SAS Insti- 
tute Inc. 1990) was used to evaluate the effect prey 
availability and vegetation structure had on Coyote 
visitation rates. Six independent variables were used 
in the model: the abundances of the three main prey 
genera, (1) Microtus, (2) Peromyscus, and (3) Sylvi- 
lagus, and the three vegetation variables, (4) vertical 
foliage density, (5) percent ground cover, and (6) can- 
opy cover. Each observation in the multiple regres- 
sion represented a single measure of the independent 
variable in one of the seven habitats during one of 15 
months. Thus, there were 105 observations for each 
independent variable, or a total of 630 data points for 
the regression analysis. Data were logarithmic-trans- 
formed prior to analysis to help satisfy assumption of 
normality of the residuals. Collinearity diagnostics, 
such as tolerance values and variance inflation factors, 
indicated a lack of colinearity between the regres- 
sors, hence all independent variables were retained in 
the model. 

Simple linear regression, performed with SAS PROC 
REG (SAS Institute Inc. 1990), was used to compare 
Coyote visitation rates against mice abundances in 
each of the two adjacent habitats, Tallgrass Prairie I 
and Oak Woods I. We performed a separate analysis for 
the response of Coyotes to prey increase following 
manipulation of the prey’s food supply. Visitation rates 
of Coyotes around the experimental plots in Tallgrass 
Prairie II were analyzed using a one-way repeated 
measures analysis of variance ((mANOVA), with SAS 
PROC GLM (SAS Institute Inc. 1990). Inferences 
were based on Huynh-Feldt adjusted P-values for the 
rmANOVA. 


Results 
Among-habitat prey fluctuations 

There was variation in spatial-temporal abundances 
of prey species (Figure 2) with significant species 
(R? = 0.658, Fro. 264) = 6.21, P = 0.002), time (Fry 264) = 
21.29, P < 0.001), and time? (Fy, 264) = 14.17, P < 
0.001) effects, indicating that overall numbers of the 
three prey differed significantly. The significant time 
X species (Fy 264) = 18.29, P < 0.001) and time* x spe- 
cies (Fi, 964; = 11.74, P < 0.001) interactions indi- 
cated that the rate at which prey species numbers 
changed also differed significantly. This was primar- 
ily due to the substantial increase in mice numbers 
(Figure 2). Furthermore, the significant time x habi- 
tat x species (Fry. 264) = 4.51, P < 0.001) and time? x 
habitat x species (Fi, 364; = 4-84, P < 0.001) inter- 
actions indicated that the differential rates in changes 
of numbers of species also changed among the dif- 
ferent habitats. For example, the highest abundances 
of mice and voles occurred in November 1994 in the 
habitats Oak Woods I and Tallgrass Prairie I, res- 
pectively. The highest Sylvilagus abundance was docu- 
mented in October 1994 in the Oldfield, with nearly 
equally high numbers recorded in July 1994 in the 


RANDA and YUNGER: SCENT STATION VISITATION RATES OF COYOTES 345 


two prairie sites, probably following the weaning period 
of young rabbits in northern Illinois (Hoffmeister 1989). 
These abundance peaks are reflected in the quadratic 
function of the change in prey numbers over time. 
Among-habitat Coyote visitation rates 

Scent station visitation rates fluctuated widely among 
habitats and over time; no signs of Coyotes were found 
in Oak Woods II and the majority of tracks were re- 
corded in the grassland habitats (Figure 3). Overall, 
the multiple regression of Coyote scent station visita- 
tion rates across Fermilab with the three main prey 
(Figure 4) and the three habitat structure variables 
(Figure 5) was significant (R* = 0.241, Fy, og) = 4.40, 
P = (0.003; Table 1). The unexplained variance in the 
model may be attributed to the observed variation in 
Coyote visitation (Figure 3) and 0 values recorded for 
prey abundances. Based upon results of the multiple 
regression, we conducted a separate analysis to discern 
whether the visitation rates in grasslands by Coyotes 
was due to prey (Microtus) or avoidance of wooded 
areas, from which voles were absent. Regression of 
scent station visitation rates with vole abundance, ex- 
cluding data from the two oak woods, was not signi- 
ficant (7? = 0.050, Fy, 73; = 3.75, P = 0.057). 

In comparing Coyote visitation in the two adjacent 
habitats, Tallgrass Prairie I and Oak Woods I, scent 
station visitation rates, when averaged over the study 
period, were over three times greater in the prairie 
(0.035 + 0.011 (mean + SE ); range 0.000—0.0645), 
than the woods (0.008, + 0.003, range 0.000—0.0179; 
tog) = 2-486, P = 0.026). However, mice abundance 
was two times greater in the oak woods (4.7 individ- 
uals per transect + 1.4, range 0.0—19.3) than the 
prairie (2.3 individuals per transect + 0.6, range 
0.3—8.0; fig) = 1.637, P = 0.113). 


Within-habitat Coyote visitation rates 

The fire in Tallgrass Prairie I did not result in a 
substantial increase in prey (Figure 2). However, the 
perturbation in Oak Woods I was followed by a con- 
siderable increase in mice abundance, enabling a 
within-habitat comparison to Coyote activity. Despite 
the high prey abundance, Coyote visitation rates were 
not significantly related to mice abundance (7° = 0.158, 
Fry, 13) = 1.03, P = 0.389) in Oak Woods I. 

Food supplementation in Tallgrass Prairie II resulted 
in an approximately three-fold increase in numbers 
of mice compared to the non-supplemented plots by 
February 1995 (Figure 6). Numbers subsequently dec- 
lined until densities converged on the supplemented 
and non-supplemented plots in May 1995. Coyotes 
responded to the peak in mice densities with mean 
visitation around the food supplemented plots over five 
times greater than on non-supplemented plots in Feb- 
ruary 1995 (Figure 6). Coyote visitation around these 
plots in the ensuing three months reflected the trend 
in prey availability. The result was a significant food 
effect (Fi, >, = 25.60, P = 0.037), a significant time 
effect (Fi; 6 = 9-79, P = 0.013), and a significant 


346 THE CANADIAN FIELD-NATURALIST Vol. 118 


@—® Tallgrass Prairie I &—L Oak Woods II 
4— A Oldfield 


O--O Shrubby Oldfield 
v--v Brome Grass Field 


C—O Tallgrass Prairie II 


@— 8 Oak Woods I 


Peromyscus 


Microtus 


Mean Number of Individuals per 200-m Transect 


Months 


FiGuRE 2. Abundances of the three main prey, Peromyscus, Microtus, and Sylvilagus, in each of the seven habitats over time. 


2004 


0.35 


0.30 


O25 


0.20 


0.15 


0.10 


0.05 


Mean Scent Station Visitation Rate 
(proportion stations visited / operative days) 


0.00 


RANDA and YUNGER: SCENT STATION VISITATION RATES OF COYOTES 347 


Tallgrass Prairie I 


Tallgrass Prairie II 
Oakwoods I 


Oakwoods II 
Oldfield 

Shrubby Oldfield 
Brome Grass Field 


Months 


FicureE 3. Habitat visitation of Coyotes, based upon monthly average scent station visitation rates, over time among the 


seven habitats. 


visitation X time interaction (F/; ) = 10.32, P = 0.012). 


Vegetation characteristics of habitats 

Of the three vegetation metrics collected, percent 
ground cover and canopy cover estimates were fairly 
consistent between the non-growing and growing 
seasons for each of the seven habitats sampled (Table 
2). Vertical foliage density was greater in the growing 
season than the non-growing season for all habitats, 
reflecting the increase in above-ground plant biomass. 
Tallgrass Prairie I had the greatest difference in ground 
cover and foliage density between these seasons. This 
was due to the almost complete absence of litter and 
standing vegetation during the March survey, except 
for small clumps of tall grasses, a result of the previous 
fall’s fire. There were similar trends of vegetation met- 
rics among similar habitats. Vertical foliage density 
was greatest in the prairies compared to other habitats 
during the growing season, but was relatively low for 
the woods. The habitats dominated by grasses, the 
prairies and brome grass field, exhibited > 97% ground 
cover, exceeding estimates of other habitats. Canopy 
cover, as expected, was highest in the woods, with 
Oakwoods II having a denser overstory than Oak- 
woods I. The presence of canopy in the brome grass 


field was attributed to a few isolated trees located 
near the middle of the field, which was not expansive 
enough to markedly influence Coyote visitation. 


Discussion 

Coyote habitat visitation, measured through scent 
station visitation rates, fluctuated greatly over time, 
showing no clear temporal pattern among habitats. 
However, overall visitation indicated an extensive use 
of grasslands and avoidance of woodlands. Through- 
out the duration of the study, there was no evidence 
of Coyote tracks or scat in Oak Woods II, a mature 
woodlot with few gaps in its overstory, even though 
its entire western side bordered a tallgrass prairie, a 
habitat where Coyote scat and tracks in snow were 
observed. Oak Woods I also experienced relatively 
little Coyote visitation during most of the study. 
Although scent station transects were 50 m from 
habitat edges, all Coyote tracks were observed at the 
western end of the transects, which were closest to a 
grassy area near the woods. The shrubby oldfield 
experienced slightly more visitation, overall, than the 
woods, but tracks were found primarily at scent sta- 
tions within open grassy areas and in the western por- 


348 THE CANADIAN FIELD-NATURALIST Vol. 118 


Peromyscus 


Sylvilagus 


Mean Coyote Visitation Rate (proportion stations visited / operative days) 


0.0 0.5 1.0 IS 2.0 De) 3.0 


Mean Number of Prey per 200-m Transect 


Ficure 4. Partial regression plots for activity of Coyotes vs. abundances of each of the three main prey species. Each point 
represents a single observation recorded during | of 15 months in one of the seven habitats. 


) 2004 RANDA and YUNGER: SCENT STATION VISITATION RATES OF COYOTES 349 


vertical foliage density 


canopy cover 


ground cover 


Mean Coyote Vistation Rate (proportion stations visited / operative nights) 


Means of Vegetation Variables 


| Ficure 5. Partial regression plots for activity of Coyotes vs. each of the three vegetation variables. Each point represents a 
single observation recorded during | of 15 months in one of the seven habitats. 


350 


GY 


WSS 


WN 


4 
‘ 


Lf 


Ny 
NS 


SS 


fee 
KK 


Mean Number of Peromyscus per Plot 


Yj 
Y 
j 
L 
Y 


Months 


THE CANADIAN FIELD-NATURALIST 


—@— Peromyscus density, +F 
—O©Q-—. Peromyscus density, -F 


WZ, Visitation rate, +F 
[___] Visitation rate, -F 


Vol. 118 


Mean Coyote Visitation Rate 
(proportion stations visited / operative nights) 


FIGURE 6. Comparison of activity of Coyotes and abundance of mice on food-supplemented (+F) and non-supplemented (-F) 


plots in Tallgrass Prairie II. 


tion of this habitat which bordered a tallgrass prairie. 
These observations were supported by the significant 
negative relationship of canopy cover to Coyote visita- 
tion rates in the multiple regression analysis. Hence, 
Coyotes may move a limited distance into woods but 
tend to avoid core areas of heavily wooded habitats. 
This behavior was corroborated by a two-year radio 
telemetry study of Coyotes in northeastern Illinois 
(Roth et al. 1999°). 

The multiple regression analysis also indicated scent 
station visitation rates by Coyotes were affected by 
abundance of voles, which was not found in the oak 
woods studied. However, a simple regression of Coy- 


TABLE |. Parameter estimates for independent variables used 
in the multiple regression analysis of Fermilab Coyote activities. 


Variable df Regression 
coefficient Jz 

Intercept 0.0261 0.040 
Peromyscus 1 -0.0082 0.192 
Microtus 1 0.1192 0.006 
Sylvilagus -0.0169 0.359 
Foliage Density 1 -0.0441 0.295 
Canopy Cover 1 -0.0632 0.004 
Ground Cover 1 0.0082 0.871 


ote visitation rates on vole abundance, excluding data 
from the two woods, did not reveal a significant rela- 
tionship. Thus, it appeared that factors other than prey, 
such as vegetation structure, strongly influenced Coy- 
ote habitat visitation at Fermilab. In addition, overall 
visitation rates were not related to the abundance of 
Eastern Cottontails, which comprised the greatest 
biomass of prey in the Coyote diets (Randa 1996). 
This could possibly be attributed to the association of 
cottontail rabbits along habitat edges (Althoff et al. 
1997; Mankin and Warner 1999; Smith and Livaitis 
2000). Although some of the scent station transects 
were located within habitat areas where rabbits were 
found, Coyotes could have spent more time foraging 
for rabbits along these edges where transects were 
not located. 

The selective use or visitation of grasslands by the 
Coyotes in this investigation may be associated with 
their historical range. Prior to European settlement, 
Coyotes occupied the southwest and central plains 
of the U. S. including grassland of southern Canada. 
Clearing of forests and extermination of the Grey Wolf 
(Canis lupus) facilitated the north- and eastward expan- 
sion of Coyotes. Coyotes moved northward along the 
Great Lakes, and colonized the northeastern U.S. 


) 


2004. RANDA and YUNGER: SCENT STATION VISITATION RATES OF COYOTES 35] 
TABLE 2. Summary of vegetation characteristics for the seven different habitats sampled twice during 1995 at Fermilab; NG 
= non-growing season sampling, G = growing season sampling. 
Mean Vertical Mean Percent Mean Canopy 

Foliage Density* Ground Cover Cover? 
Habitat NG G NG G NG G 
Tallgrass Prairie I 0.031 0.703 352, 98.0 0.000 0.000 
Tallgrass Prairie II 0.320 0.761 OED 97.5 0.000 0.000 
Oakwoods I 0.196 0.377 94.7 89.7 0.750 0.792 
Oakwoods II 0.132 0.172 66.4 52.4 1.000 1.000 
Oldfield 0.328 0.459 89.4 89.2 0.000 0.000 
Shrubby Oldfield 0.248 0.587 96.7 89.5 0.389 0.340 
Brome Grass Field 0.180 0.346 100.0 99.7 0.042 0.021 


“Based upon the average proportion of three profile boards obscured by vegetation for 24 sampling stations per habitat. 
>Based upon presence/absence scores (1/0 respectively) for 24 sampling stations per habitat. 


(Richens and Hugie 1974) and the Midwest (Moore 
and Parker 1992) by the 1940s. Most Coyotes still in- 
habit deserts and grasslands, and those in regions with 
forests tend to use open, non-forested areas in a great- 
er proportion than available (Todd et al. 1981; Toweill 
and Anthony 1988; Cypher 1991; Holzman et al. 
1992; Murray et al. 1994; Kamler and Gipson 2000). 
Tracks of Coyotes observed in the snow at our study 
site indicated that grasslands, such as the brome grass 
field and tallgrass prairie, were occasionally traversed 
by Coyote tracks following a relatively straight path, 
reflecting non-foraging behavior. Hence, these open 
areas may not only serve as preferential foraging sites 
but as areas facilitating movement between habitats. 
A preferential use of grasslands may explain why 
chipmunks and tree squirrels, which were moderately 
common in the woodlands, were rarely detected in 
Coyote diets (Randa 1996). 

Selective use of grasslands may have restricted the 
ability of Coyotes to respond spatially and temporally 
to localized patches of abundant prey within a habi- 
tat. In Fall 1994, when mice densities in Oak Woods 
I were more than twice the prey abundances at any 
other location, Coyote visitation did not increase sig- 
nificantly in that habitat. This discrepancy of prey abun- 
dance and coyote response in woodland corresponds 
with findings in southeastern Quebec (Richer et al. 
2002). However, Coyotes demonstrated an ability to 
closely track prey within a grassland (Tallgrass Prairie 
II) as their visitation rates were correlated with ex- 
perimentally manipulated abundant patches of mice. 
The high level and subsequent decline in Coyote visi- 
tation rates detected on scent stations around food- 
manipulated plots paralleled the change in mice 
densities. Hence, once mice decreased following a 
peak in densities on these plots, Coyote visitation rates 
likewise diminished. It was unlikely that Coyotes were 
responding to a novel item in the environment (such 
as the odor of the food) because supplementation had 
been initiated four months prior to the scent station 
monitoring. Because voles were not detected in this 


habitat during the experiment, a similar test of within- 
habitat Coyote response to abundant patches of this 
commonly consumed prey was not feasible. Yet, as 
with mice, Coyotes could have responded to isolated 
patches of voles within habitats throughout Fermilab, 
given that voles were still a major component of Coy- 
ote diets despite their low availability. 

One limitation of the scent station transects was the 
difficulty in identifying Coyote responses to possible 
small, isolated patches of prey. Also, due to the large 
areas over which Coyotes may move, there was not 
sufficient separation to avoid visitation by the same 
individual between scent station transects (Diefenbach 
et al. 1994; Roughton and Sweeny 1982). However, 
by intensively sampling each habitat, these transects 
were effective in discerning overall trends (i.e., among- 
habitat comparisons) in Coyote visitation rates and rel- 
ative habitat use; they were not intended to indicate 
finer-scale or localized responses (e.g., patch-level 
foraging) within the study site. 

The results we obtained from both the experimen- 
tal plots and scent station transects demonstrated that 
Coyotes exhibit variable responses to prey availability, 
depending upon habitat type. Reacting relatively quick- 
ly to a local increase in prey, as seen around the ex- 
perimental plots in prairie, would enhance Coyote 
foraging efficiency. However, preferential use of open 
areas, as suggested by the transect data, may con- 
strain foraging efficiency during periods of very low 
prey densities when alternative prey in woodlands is 
under-utilized. Hence, supporting Coyote populations 
near urbanized areas, as in northern Illinois, requires 
open land expanses such as grasslands to facilitate 
movements across large spatial extents within the land- 
scape and to provide an adequate prey base. 


Acknowledgments 

P. L. Meserve lent support and guidance throughout 
the course of this investigation. S. Scheiner and C. 
von Ende provided statistical advice. D. Cooper shared 
in much of the field activity. J. Mendelson, E. Miller, 


392 


C. Petersen, B. Satterfield, and several anonymous 
reviewers provided helpful comments on earlier drafts 
of this manuscript. R. Walton oversaw Fermilab activ- 
ities associated with this project and helped obtain 
funding. The National Environmental Research Park 
of the U.S. Department of Energy (MO-RS023), and 
the Department of Biological Sciences, Northern 
Illinois University provided funding for this study. 


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Received 27 February 2003 
Accepted 14 October 2004 


Presence of Cavities in Snags Retained in Forest Cutblocks: 
Do Management Policies Promote Species Retention? 


Ki T. Everett!:? AND KEN A. OTTER? 


‘Forestry Program, University of Northern British Columbia, Prince George, British Columbia V2N 4Z9 Canada 

"Ecosystem Science & Management Program, University of Northern British Columbia, Prince George, British Columbia 
V2N 4Z9 Canada 

3Current Address: Department of Biology, University of Victoria, Victoria, British Columbia V8W 2Y2 Canada 


Everett, Kim T., and Ken A. Otter. 2004. Presence of cavities in snags retained in forest cutblocks: do management policies 
promote species retention? Canadian Field-Naturalist 118(3): 354-359. 


Tree cavities, which are frequently excavated by primary cavity nesters, are typically used by a number of avian and non-avian 
species and are thus important components in maintaining biodiversity in forest ecosystems. One way to provide these 
habitat opportunities in harvested areas is through the retention of snags. In this study, we assessed the habitat and snag 
characteristics that promote cavity excavation, using the presence of cavities to infer activity of primary cavity excavators. 
Snags retained closer to the forest/cutblock edge contained a greater density of cavities than trees further from edge. However, 
the proportion of cavities found within cutblocks declined at a more rapid rate with distance from edge than did those in adja- 
cent forested stands. There was also a tendency for cavities to occur more frequently in trees that were at the advanced stages 
of decay. The results of our study suggest management for snags in harvest areas should include the retention of snags 


closer to the forest edge combined with incorporating trees showing signs of advanced decay. 


Key Words: tree cavities, snags, forest edge, birds, mammals, reptiles, British Columbia. 


Snags (standing dead trees) provide specialized hab- 
itat for many species, which in turn play an integral role 
in maintaining ecosystem functions (Machmer and 
Steeger 1995*). Approximately 16% of the indigenous 
birds, mammals, and reptiles in British Columbia de- 
pend to some extent on snags or cavities for repro- 
duction, feeding, and shelter (Backhouse and Lousier 
1991*). These dependent species then create habitat 
opportunities for other wildlife species, facilitate seed 
dispersal, increase soil aeration, accelerate organic 
decomposition, and can even aid in controlling forest 
pests (Machmer and Steeger 1995*). Although snags 
play an important role in the ecosystem, their removal 
has traditionally occurred in past forest management 
(ostensibly to reduce hazards — lightning attraction, 
falling snags, etc. — and foci of infection for healthy 
trees). 

By retaining trees of varying ages, diameters, and 
decay classes in cutblocks, the vertical structure of 
vegetation, total percent vegetation, and percent decid- 
uous cover increase. These changes can lead to greater 
species richness and diversity in both plant and mam- 
mal communities (Davis et al. 1999*). Population den- 
sities of cavity-nesting species are often limited by 
potential nest sites (Runde and Capen 1987), and inten- 
sive forest management that removes snags reduces 
their densities in managed stands compared to natural 
stands (Haapanen 1965; Pojar 1995). 

Primary cavity nesters (those which excavate their 
own cavities, such as woodpeckers and some chicka- 
dees: Thomas et al. 1979) appear to prefer stands with 
a high number of snags (Zarnowitz and Manuwal 1985; 


Raphael and White 1984). As most primary cavity- 
nesting birds rarely occupy nests that they themselves 
have not excavated (Short 1979), suitable trees, in- 
cluding snags, must be available for excavation for 
nests or roosts. Snags offer softer excavating substrates 
as well as important foraging sites for these species. 
Conversion of natural forests into intensively managed 
stands which lack snags may, therefore, have long-term 
negative effects on the entire cavity-nesting populations, 
as secondary cavity-nesting species rely on primary 
excavators for cavity creation (Peterson and Grubb 
1983). 

Bodies that govern forestry practices recognize the 
importance of snags, and advocate the retention of a 
variety of tree species at various stages of decay in 
managed forests (e.g., Machmer and Steeger 1995*). 
In an effort to reduce the impact of large openings 
created by cutblocks, some forest companies retain 
windfirm deciduous trees and advanced regeneration 
that fall below utilization standards to aid in main- 
taining structural diversity in cutblocks. This limits 
the distance wildlife must travel through the cutblock 
without vegetative cover. The traditional paradigm has 
been that efforts such as these will be sufficient to 
maintain wildlife and biodiversity throughout managed 
forests. 

One aspect of snag retention policy that has been 
overlooked, however, is the behavioral characteristics 
of primary cavity-nesting species that may deter mem- 
bers of this guild from occupying trees left specifically 
for their use. Many forest birds avoid areas outside 
forested cover (Hegner 1985; Desrochers and Hannon 


354 


2004 


1997), as open spaces leave these species susceptible 
to hawk and falcon predation (Corral 1989). Several 
studies have shown that many forest dwelling species 
are reluctant to cross gaps between fragmented forest- 
ed areas (Lens and Dhondt 1994; Desrochers and 
Hannon 1997). Desrochers and Hannon (1997) found 
that forest birds were three times less likely to cross 
gaps 70 metres wide, and eight times less likely to 
cross gaps 100 metres wide compared to similar dis- 
tances within the forest. Machtans et al. (1996) also 
found that movement rates of forest birds across cut- 
blocks were significantly lower than movement rates 
within the forest. Instead, forest birds preferred to travel 
much greater distances through corridors, rather than 
crossing narrow gaps (Desrochers and Hannon 1997). 

The avoidance of open spaces indicated that forest 
birds — with the majority of those studied belonging 
to the cavity-nesting guild — have fewer habitat oppor- 
tunities as the landscape becomes increasingly frag- 
mented. Although retained snags may be scattered 
throughout cutblocks to provide nest sites, the gap 
between the forest edge and the snag may be too sub- 
stantial for birds to cross. In turn, the number of cavity- 
nesting birds may decline with increased distance into 
cutblocks as compared to the surrounding forest. 

In this study, we documented the presence of cavi- 
ties in retained snags within forest cutblocks as a mea- 
sure of use by cavity-excavating species, and compared 
this to similar trees found within the surrounding for- 
est. If primary cavity nesting species avoid openings 
in forest cover, we predict fewer cavities as one travels 
further from the forest edge into cutblocks. By assess- 
ing attributes of cavity-possessing trees, we attempt 
to identify which tree characteristics and tree species 
cavity nesting birds tend to utilize. Our aim is to 
provide well-substantiated recommendations towards 
the placement and creation of retained snags within 
cutblocks that maximize potential usage by cavity 
nesting species. 


Methods 
Study Area 

This study was carried out from 30 September to 5 
November 2000, approximately 30 kilometers west of 
Prince George, British Columbia (53°36'35"N, 
122°57'29"W). All samples were located within the 
Sub-Boreal Spruce biogeoclimatic zone, which covers 
much of the central interior of British Columbia 
(MacKinnon et al. 1992). The study area was primarily 
composed of coniferous and mixedwood forests, with 
species including Lodgepole Pine (Pinus contorta 
var. latifolia), hybrid White Spruce (Picea glauca x 
engelmannii), Sub-alpine Fir (Abies lasiocarpa), Doug- 
las-fir (Pseudotsuga menziesii var. glauca), Trembling 
Aspen (Populus tremuloides), Black Cottonwood (Pop- 
ulus balsamifera ssp. trichocarpa) and Paper Birch 
(Betula papyrifera). Harvested areas are typically re- 
planted with Lodgepole Pine, with a small component 
of hybrid White Spruce on moister sites. 


EVERETT AND OTTER: PRESENCE OF CAVITIES IN SNAGS 355 


Sampling Procedure 

We randomly selected eight cutblocks for the survey 
from a Forest Development Map. The cutblocks ranged 
in age from 3 to 10 years post-harvest, and from 30 to 
60 hectares in size. A line-transect method of sampling 
was used. In this method, all retained snags falling with- 
in 5 metres of each side of the transect were measured. 
We ran transects on a bearing perpendicular to the for- 
est edge, with a random point of commencement along 
the edge being selected from the Forest Development 
map. Measurements were carried out until the centre- 
line of the cutblock was reached and at least 20 snags 
were encountered. In one case, the cutblock contained 
only 17 retained snags. Due to the minimal number 
of retention trees present, the line-transect method 
would have been relatively ineffective; therefore; all 
retention trees in the cutblock were measured. 

Snags were defined as any dead, standing tree (Smith 
et al. 1997: 489). We recorded all cavities that were 
either actively excavated or were natural cavities of 
similar size, which displayed signs of active use (e.g., 
modification to the cavity, presence of feathers in 
entrance, etc.). Cavities were only recorded if they had 
a minimum diameter of 3.5 cm, as the cavity had to 
be large enough for use by a small bird or mammal. 
However, as our study was looking for evidence of use 
of the trees rather than specifically focusing on nesting, 
we did not distinguish between cavities used for roost- 
ing versus those where nesting had occurred. For each 
snag encountered along the transect, we recorded sev- 
eral attributes: tree species, height, diameter at breast 
height (DBH, measured at 1.3 m from the base of the 
tree), decay class, number of cavities, distance from 
the nearest timber edge, percent overhead cover, and 
the number of snags and live trees within an 11.28 m 
radius around the trees. Only trees greater than 10 cen- 
timeters DBH were recorded, as this reduced the chance 
of incorporating regeneration into the study. Decay 
class was determined using a six-step hardwood classi- 
fication scheme combined with a corresponding nine- 
step coniferous classification scheme (Ministry of 
Forests 2000*). 

Similar transects were placed in the adjacent forest- 
ed stands running perpendicular to the edge. In order 
to measure a similar number of trees over the same dis- 
tance in both habitat types, every third tree classified 
as decay class 2 or greater was measured. In order to 
determine the initial tree for measurement along each 
transect, a coin toss was completed to decide if the 
first or second tree should be measured. For each tree 
encountered along transects in forested stands, the 
same attributes were measured as those collected for 
cutblock sampling. 


Statistical Analysis 

A multiple regression was used to determine if the 
habitat (forest or cutblock) and distance from the for- 
est edge influenced the proportion of cavities found 
(SYSTAT 9.01). To compare the distribution of tree 


356 


types across the two habitats, we employed a Chi- 
square and used a Mann-Whitney U test to compare 
the frequency of decay classes of trees between cut- 
blocks and forested stands. 

To determine which characteristics tended to predict 
the presence of cavities in wildlife trees, we conduct- 
ed a Principal Component Analysis (PCA) on trees in 
the forested stands. As the forest stands constitute the 
natural habitat of many of the primary cavity-nesting 
species, determining the characteristics of cavity-bear- 
ing trees in these areas may give insight into which 
types of wildlife trees are important to retain in 
cutblocks. The following characteristics of the focal 
tree and the immediate area were subjected to PCA: 
height, diameter, and decay class of the focal tree, along 
with the total number of trees, number of live trees, 
number of snags, and percent overhead cover within 
the 11.28 metre radius plot. A linear regression was 
used to compare the number of tree cavities against 
the PCA factor scores. The PCA was carried out in 
SYSTAT 9.01 using Varimax rotation. 

Two retention trees measured had extremely high 
numbers of cavities and were identified during initial 
analysis as significant outliers in the data set (Durbin- 
Watson D Statistic = 1.984, P < 0.01). These were 
excluded from subsequent analysis. 


Results 

In total, 185 trees were measured at our eight sam- 
pling sites; 94 trees within cutblocks and 91 trees within 
forested stands. We found 46 cavity-bearing trees, con- 
taining a total of 67 cavities. Of the 46 trees containing 
cavities, 29 were deciduous while the remaining 17 
were conifers. 

A greater proportion of deciduous trees contained 
cavities compared to coniferous species. Species of 
willow had the highest proportion of cavities, with 
two of the three trees observed containing cavities. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Trembling aspen was found to have the second highest 
frequency of cavities with 17 of the 62 trees observed 
containing cavities. A breakdown of the number of 
cavities recorded by species is given in Table 1. 

The species composition of trees differed between 
the two habitats (x? = 76.6, df = 6, p< 0.01 — Figure 1). 
Snags in cutblocks were primarily deciduous species; 
although these trees were also present in forested 
stands, the latter stands had a proportionately higher 
coniferous component. 


Distance From Edge and Cavity Abundance 

Distance from forest edge and stand type (cutblock 
or forest) both had a significant effect on the number 
of cavities found (multiple regression: 17=0.051, 
Fo 19) = 4-857, p = 0.009). The further a tree was 
located from the forest-cutblock edge, the lower the 
frequency of cavities (p = 0.027). There was also a sig- 
nificant effect between the stand types, as more cavities 
were found in the forest than were found in the cut- 
block (p = 0.020). In the forest stands, the decline in 
number of cavities found with distance from the edge 
was not as dramatic as that seen in cutblocks (Figure 
2a and b). 


Stand Characteristics Associated with Snags 

The principal component analysis identified two 
factors that accounted for 56.2% of the total variance 
among retained snags (Table 2). Principal component 
one (PC 1) accounted for 31.2% of the variance, with 
strong positive weightings for a high number of total 
trees and a high number of live trees within an 11.3 m 
radius plot. As a result, PC 1 can be considered a meas- 
ure of the density of trees surrounding the snag. 

Principal component two (PC 2) accounted for 
25.0% of the total variance, with a strong positive weight- 
ing for decay class and a strong negative weighting 
for tree height. PC 2 indicates trees at the latter stages 
of the decay process, as tree height frequently decreases 


TABLE |. A breakdown of the number of cavities by tree species. 


Tree Species Number 
of trees 
observed 
Trembling Aspen 
Populus tremuloides 62 
Lodgepole Pine 
Pinus contorta var. latifolia 52 
Paper Birch 
Betula papyrifera Sy) 
Hybrid Spruce 
Picea glauca X engelmannii 24 
Willow 
Salix sp. 3 
Black Cottonwood 
Populus balsamifera ssp. trichocarpa 8 
Douglas Fir 
Pseudotsuga menziesii glauca 4 


Total (Average) 185 


Total number 
of trees 


Percentage 


of trees Total number 


with cavities with cavities of cavities 
17 27.4 32 
13 25.0 14 
8 25.0 10 
4 16.7 5 
2 66.7 3 
2 25.0 3) 
0 0 0 
46 (24.9) 67 


2004 
70 | Cutblock 
” Forest 
a) 
o 
Qa 
2) 
S38 
ae 
ES 
25 


CJ 
Q 
n 
x 
o 
” 


Populus balsamifera 
Populus tremuloides 
Betula papyrifera 
Pseudotsuga menziesii 

Pinus contorta 

Picea glauca 


Tree Species 


FiGure 1. Distribution of sampled snags within cutblocks 
versus surrounding forest stands. 


as the tree reaches the higher decay classes (Towers 
et al. 1992). 


Response of Cavity Nesters to Stand Characteristics 

There was no significant relationship between the 
number of cavities found and PC | (regression: r* = 
0.003, Fo 99) = 0.281, p = 0.60). There was, however, a 
significant effect between the number of cavities being 
found and PC 2 (regression: 1? = 0.092, F,, 99) = 9.13, 
p = 0.003) indicating that more cavities occurred in 
trees with a higher amount of decay. 

On average, the snags retained in clearcuts had 
less decay than the comparison cohort of trees in the 
forested areas (MWU: U = 2918.5, n= 93, 91, p<0.01). 


Table 2. Principal component analysis of stand characteris- 
tics in forest stands. The first two Principal Components ac- 
count for greater than 50% of the total variance among stand 
attributes; therefore only these two factors were considered 
in subsequent analysis. Variables that were within 10% of 
the highest loading were considered to be strong weightings, 
and are shown in bold. 


Variable Real PE2 
(clustering of (decay 
retention trees) stage) 
Tree Diameter -0.407 -0.395 
Tree Height 0.272 -0.714 
Decay Class -0.353 0.738 
Total Number of Trees 0.947 0.212 
Number of Live Trees 0.956 0.115 
Number of Snags 0.000 0.392 
Percent Overhead Cover -0.029 0.573 
Percent of Total Variance 
explained by each Factor SIRS 25.02 


EVERETT AND OTTER: PRESENCE OF CAVITIES IN SNAGS 


a. Forest 


n 
ut 
= 
> 
<x 
oO 
LL 
Oo 
a 0 100 200 300 400 
rea) 
=| b. Cutblock 
=) 
Zz 
[o) 
TDC O 
yea ae I EE Be UE 
100 200 300 400 


DISTANCE FROM EDGE (m) 


FIGURE 2. Regressions of distance from forest edge against the 
number of cavities located in trees in forested stands 
(a — shown by a dotted line) and cutblocks (b — shown 
by dashed line). 


Discussion 
Edge Effects on Cavity-nesting Birds 

The number of cavities located within both cutblocks 
and forested stands were related to the distance from 
forest edge. Trees located farther from the forest edge 
had a tendency to contain fewer cavities than trees 
located closer to the forest edge. There was also a 
greater abundance of cavities located within forested 
stands than within cutblocks. Yet, although the number 
of cavities declined rapidly as the distance from cut- 
block edge increased, the equivalent decline was less 
rapid in forested stands. This suggests that retained 
snags farther from forest cover were underutilized by 
cavity-excavating species. 

One possible bias in our results might arise if cavi- 
ties found in the cutblocks existed prior to harvesting. 
Although we focused on measuring cavities that ap- 
peared to be recently used/excavated, it was occasion- 
ally difficult to distinguish between recently excavated 
cavities and cavities that may have existed prior to 
logging. However, if some of the cavities existed prior 
to harvest our results would over-estimate the level of 
activity of cavity-excavating species in cutblocks, thus 
minimizing the perceived effect of harvesting on cavity 
nesters. Despite the possible bias of including pre-exist- 
ing cavities, we continued to see a reduced number of 


358 


cavities in clearcuts. Thus, the magnitude of the effect 
we report may be even more pronounced between the 
different habitats. 

Several possible explanations may exist for the low- 
er densities of cavities within cutblocks. The reduced 
number of cavities in cutblock trees could be due to 
the reluctance of many species of forest birds to cross 
large openings. Desrochers and Hannon (1997) found 
that forest birds were extremely vulnerable to raptors 
when they fly in the open, and thus were hesitant to 
enter such areas. Secondly, the lower densities of tree 
cavities within cutblocks could be due to the retention 
of trees with minimal habitat value for cavity nesting 
species. Cavity nesters tended to prefer deciduous tree 
species, as a greater number of cavities were found in 
these trees than in coniferous tree species. In conifer- 
dominated ecosystems, such as boreal and sub-boreal 
forests, deciduous trees often have greater heartwood 
decay than conifers and may be preferred as nesting 
substrate once the trees become snags, resulting in 
increased avian diversity (Martin and Eadie 1999). 
Yet, deciduous species accounted for a larger number 
of snags in the cutblocks than the forests in our study, 
and there was still a lower number of cavities in cut- 
blocks than neighboring forests. While the retention 
of these deciduous snags likely increases diversity 
along the edge, the reluctance of birds to venture into 
cutblocks may have a larger influence on snag use 
farther from forest cover. 

The highest density of cavities was located within 
100 metres of the forest edge. Higher bird populations 
located at the forest edge compared to the community 
interiors has also been found in several other studies 
(see Gates and Gysel 1978; McElveen 1979; and 
Streikle and Dickson 1980). Several theories exist to 
support such behaviour. Marcot (1983) suggested that 
many cavity-nesting species nest in the nearby forest, 
whereas they use cutblocks for foraging, resulting in 
higher densities of birds seen along the community 
edge. Both the Northern Flicker (Colaptes auratus) and 
the Downy Woodpecker (Picoides pubescens), primary 
cavity excavators, prefer to nest in old trees near open- 
ings or on the edge of the forest (Ehrlich et al. 1988). 
Our results suggest this pattern may occur within our 
study area. Cavity density was highest at edges, but 
remained high for greater distances into forests than 
into cutblocks. Thus, marginal species attracted to edges 
may prefer to either nest on the edge or penetrate into 
the forest, rather than establish nests in snags retained 
in the cutblocks. Coupled with non-edge species, this 
would create the pattern of greater cavity abundance 
in forested sites. 


Tree Attributes and Cavity Abundance 

Within forested stands, cavities were found more 
often in trees that contained a combination of high lev- 
els of decay and shorter tree height. Decayed portions 
of the tree allow for easier cavity excavation for weak- 
er cavity excavators, and the bark cover retains mois- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


ture and warmth as well as providing protection from 
predators (Runde and Capen 1987). Decayed portions 
of the tree also provide valuable foraging habitat as 
they often contain high levels of beetle larvae, carpen- 
ter ants, and termite activity, all of which act as food 
resources for woodpeckers (Mannan et al. 1980). This 
too may explain the lower use of cavities in cutblocks, 
as trees retained in these areas had less decay com- 
pared to trees sampled randomly in the surrounding 
forests. This is likely the result of trees with greater 
decay being removed as potential hazards during for- 
estry operations. Retention of these higher decay class 
trees, however, may be valuable in future planning. It 
is important to retain trees at a variety of levels of 
decay, as the nesting success of some primary cavity 
excavators is higher in snags in the 2-3 decay class 
range (Fort and Otter 2004). 


Management Implications and Recommendations 

Primary cavity nesters require specific habitat con- 
ditions (Conner et al. 1976), and without these condi- 
tions territory selection and reproductive success may 
be compromised (Kilham 1966). As a result, popula- 
tion densities of both primary and secondary cavity 
nesters may decline. 

In order to maintain the population levels of pri- 
mary cavity-excavators it is essential to provide ade- 
quate habitat through efficient forest management. 
Snags and snag patches may be most efficient if they 
are placed within 100 metres of the forest edge, as 
this limits susceptibility of forest-dwelling birds to 
predators. The retention of trees for the purpose of 
managing cavity nesting birds should focus on main- 
taining a high deciduous component within the stand. 
Cavity nesters tended to select a high level of decidu- 
ous trees for cavity excavation throughout this study. 
Maintaining a variety of deciduous and coniferous 
trees within cutblocks could prove to be beneficial 
for population densities of cavity nesters, as well as 
many other species of snag users. 


Acknowledgments 

We thank Sarah Parsons, Emma Tayless, and Der- 
rick Lalonde for their help in the field, as well as Rod 
Lenton and Chris Jones from Timberline Forest Inven- 
tory Consultants Ltd. for providing maps and access 
advice. We thank Kelly Sherman for providing advice 
for sample planning as well as proof-reading numerous 
drafts. Susan Stevenson and two anonymous reviewers 
kindly provided feedback on the manuscript 


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Received 28 March 2003 
Accepted | October 2004 


Summer Diet of Two White-tailed Deer, Odocoileus virginianus, 
Populations Living at Low and High Density in Southern Québec 


CLAUDE DAIGLE!:* , MICHEL CRETE! 3, Louts LESAGE” *, JEAN-PIERRE OUELLET?, and JEAN Huor? 


' Ministére des Ressources naturelles, de la Faune et des Parcs du Québec, Direction du développement de la faune, 675 
boul. René-Lévesque est, BP 92, Québec, Québec GIR 5V7 Canada; corresponding author 

> Département de biologie et Centre d’études nordiques, Pavillon Vachon, Université Laval, Sainte-Foy, Québec G1K 7P4 
Canada 

> Département de biologie et Centre d’ études nordiques, Université du Québec a Rimouski, 300 Allée des Ursulines, Rimouski, 
Québec GS5L 3A1 Canada 

+ Present address: Environment Canada, Region de Québec, Service Canadien de la faune, 1141 Route de LEglise, C.P. 10100, 
Ste-Foy, Québec G1 V 4H5 Canada 


Daigle, Claude, Michel Créte, Louis Lesage, Jean-Pierre Ouellet, and Jean Huot. 2004. Summer diet of two White-tailed Deer, 
Odocoileus virginianus, populations living at low and high density in southern Québec. Canadian Field-Naturalist 
118(3): 360-367. 


We investigated summer diets of two White-tailed Deer (Odocoileus virginianus) populations through rumen content analyses. 
Samples from 93 deer were collected in a low density, LD (1 deer/km?) and a high density, HD (14 deer/km?) area of south- 
ern Québec during the growing seasons of 1997 and 1998. Availability of preferred forage in forests was greater in LD than 
in HD, whereas agriculture covered a larger proportion of the area in HD than LD. Rumen composition differed between the 
two populations. Deer from HD consumed less forbs and leaves of shrubs and trees than did LD deer, whereas they con- 
sumed more fruits, grasses and farm crops. The rarity in HD rumens of food items common in LD, as well as in many parts 
of the White-tailed Deer range (i.e., Lilliacae), indicated that deer could not compensate for the rarity of preferred forest 
forage by increasing foraging time and had to feed on cultivated crops. Rumen contents of LD deer had a higher level of cell 
solubles and lignin, which reflected their greater reliance on quality forage growing in forests. Feeding habits and forage 
quality can explain why deer body size decreased in HD between the 1970s and 1990s whereas LD deer remained large. 


Key Words: White-tailed Deer, Odocoileus virginianus, diet, density, forest, agriculture, plant, Québec, Canada 


Nous avons examiné le régime alimentaire estival de deux populations de cerf de Virginie (Odocoileus virginianus) par analyse 
du contenu des rumens. Des échantillons ont été prélevés sur 93 cerfs provenant d’un secteur a faible densité de cerfs, LD, 
(1 cerf/km?) et d’un secteur a haute densité, HD, (14 cerfs/km?) du sud du Québec durant la saison de croissance des végé- 
taux de 1997 et 1998. La disponibilité des aliments forestiers préférés des cerfs était plus grande dans LD que dans HD alors 
que la proportion du territoire agricole était supérieure dans HD. La composition des rumens des cerfs de chacune des régions 
différait. Les cerfs de HD ont consommé de moins grandes proportions de plantes herbacées et de feuilles d’arbustes et d’ar- 
bres que ceux du secteur LD, mais de plus grandes proportions de fruits, de graminées et de plantes agricoles. La rareté dans 
les rumens du secteur HD d’aliments communs dans ceux du secteur LD et dans plusieurs autres régions de l’aire de répar- 
tition du cerf (e.g., Lilliacae), indique que les cerfs du secteur HD ne pouvaient compenser pour la rareté de leurs aliments 
préférés par un accroissement de la durée de la quéte alimentaire, et qu’ils devaient consommer des plantes agricoles. Les 
cerfs de LD ont mangé des aliments contenant plus de solubles cellulaires et de lignine que ceux de HD, ce qui refléte une 
consommation de plantes forestieres de bonne qualité. Le régime alimentaire et la qualité de la nourriture peuvent expliquer 
pourquoi la taille des cerfs de HD a diminué entre les années 1970 et 1990 alors que celle des cerfs de LD est demeurée grande. 


White-tailed Deer, Odocoileus virginianus, are selec- 
tive feeders and although they ingest a wide variety of 
plants, only a few taxa make up large proportions of 
their diet (Johnson et al. 1995; Healy 1971; Korschgen 
et al. 1980; McCaffery et al. 1974). Plant phenology 
influences their diet; forbs and leaves of deciduous 
woody plants are common food items in spring and 
summer, and fruits become major components of the 
autumn diet. White-tailed Deer also consume mush- 
rooms in natural ecosystems (Crawford 1982; Johnson 
et al. 1995; Korschgen et al. 1980; Short 1971; Skinner 
and Telfer 1974), whereas farm crops can be major 
food sources in agricultural landscapes (Nixon et al. 
1991). During winter, White-tailed Deer switch to twigs 


of both deciduous and evergreen woody plants and, 
where snow cover allows access, they also use grasses 
and farm crops left after harvest (Johnson et al. 1995; 
McCaffery et al. 1974; McCullough 1985; Nixon et 
al. 1991; Skinner and Telfer 1974). 

Forage competition results in consumption of food 
of reduced quality or availability, which lengthens 
ingestion and digestion. At high density, forage compe- 
tition can modify feeding habits of ungulates, in partic- 
ular for small, selective species (Kie and Bowyer 1999) 
and reduce body size in cervids (Ashley et al. 1998; 
Créte et al. 1993; Hjeljord and Histol 1999; Lesage et 
al. 2001). In mid-latitude deer, forage competition 
during the growing season likely determines adult body 


360 


2004 


size because growth and replenishment of body reserves 
occur during this period of the year (Hjeljord and Histol 
1999; Lesage et al. 2001; Boucher 2004). 

On the south shore of the St. Lawrence River in 
| Québec, the carrying capacity of the deer range de- 
creases northward because suitable forest stands for 
wintering become increasingly rare with increasing 
| winter severity (Boucher 2004). Density averaged ~1 
| deer/ km? in northernmost hunting zones of Québec 
| whereas it locally exceeded 30 deer/km? in southern 
zones after populations erupted due to mild winters 
and a conservative hunting regime. Forage competi- 
_ tion in winter regulates deer numbers in the absence 
_ of Grey Wolves (Canis lupus) and under conservative 
| hunting regime (Dumont et al. 2000), but agriculture 
can help deer to survive winter by providing some food 
_ during the dormant season (Rouleau et al. 2002a). In 
recent decades, browsing pressure exhibited a pro- 
_ nounced gradient on the south shore of the St. Law- 
rence River, which resulted in a scarcity of preferred 
summer forage in southern vs. northern zones (Rou- 
| leau et al. 2002a). Vegetation sampling throughout 
| the deer range in Québec showed a general pattern of 
| negative relationship between deer density on one 
hand, and preferred forest forage and deer size on the 
_ other (Boucher 2004). Deer living at low density tend- 
ed to avoid cultivated fields (Lesage et al. 2002) and 
reached the largest size among Québec deer (Lesage 
| etal. 2001). 

We studied the summer feeding habits of White- 
| tailed Deer in two areas which differed markedly with 
respect to deer density, forage availability, impor- 
_ tance of agriculture, and winter severity. The objective 
_ of this study was to compare composition and quality 
_ of summer diets of these two White-tailed Deer pop- 
| ulations. We anticipated that preferred forage of deer 
| living at low density would be less abundant in rumens 
of deer living at high density and that the low density 
population would consume forage of higher quality 
due to relaxed forage competition during summer. 


Study Area 

Samples were collected in a low-density area (LD) 
and a high-density area (HD) (Figure 1). The LD study 
area is located in a transition zone between northern 
hardwood forests and boreal forests (Rowe 1972*). 
Among trees, Balsam Fir (Abies balsamea), White 
Spruce (Picea glauca), Eastern White Cedar (Thuja 
occidentalis), Trembling Aspen (Populus tremuloides), 
and Yellow Birch (Betula alleghaniensis) dominate 
(Lamoureux 1994*). Commercial logging has been 
intensive in this region and the spruce budworm 
(Choristoneura fumiferana) adversely impacted fir 
and spruce stands in the late 1970s. Agriculture rep- 
resents about 10% of the land use in the study area 
and hay; alfalfa (Medicago sativa), cultivated and up- 
graded pastures, oat (Avena sativa), and barley (Hor- 
deum vulgare) cover most cultivated fields. The grow- 


DAIGLE, CRETE, LESAGE, OUELLET, AND HUoT: SUMMER DIET OF DEER 36] 


ing season averages 160 days (Wilson 1971*) and 
snow cover often exceeds 50 cm for more than SO days. 
Deer density was estimated through double-count aerial 
surveys (Potvin et al. 1992) and averaged 1.1 deer/km? 
in summer 1998 (Lamoureux and Pelletier 2000*). 

Northern hardwood forests cover most of the HD 
study area. Commercial logging has affected mainly 
coniferous stands on small areas due to land owner- 
ship and forest composition (Gosselin 1994*). Most 
common tree species include Sugar Maple (Acer sac- 
charum), Red Maple (Acer rubrum), Yellow Birch, 
American Beech (Fagus grandifolia), White Ash (Frax- 
inus americana), Balsam Fir, Hemlock (Tsuga cana- 
densis), Eastern White Cedar, and White Spruce. 
Forests have been cleared for agriculture throughout 
the region. They have progressively grown back on 
abandoned farms; and introduced plants such as apple 
trees (Malus pumila) are common in today’s forests. 
Agriculture now covers about 23% of the area. Hay, 
corn (Zea mays), uncultivated pastures, cultivated 
and upgraded pastures, and alfalfa dominate cultivat- 
ed fields. The growing season averages 190 days and 
snow cover rarely exceeds 50 cm. Deer density aver- 
aged 14.1 deer/km? in summer 1996 (Dicaire 1999*); 
locally it may exceed 30 deer/km?. 


Methods 

Forage composition — We collected rumen samples 
from road kills during the growing season (13 May — 
2 November 1997 and 1998). Rumen samples were 
generally collected within 3 days after collision (maxi- 
mum 5 days), provided that identifiable forage frag- 
ments remained. Rumen content was hand mixed and 
two one-litre samples were extracted. The first litre 
was frozen for forage identification and the other pre- 
served in 4% formalin for chemical analysis. At the 
laboratory, frozen samples were thawed and washed 
through a 7.9 mm and a 4.0 mm mesh size sieve (Créte 
et al. 1990; Gauthier et al. 1989). About 50 ml of the 
particles kept by the 4.0 mm sieve were spread into a 
20 x 50 cm sampling tray (Chamrad and Box 1964) 
and covered with 1 cm of water. One hundred particles 
were systematically selected by point sampling (Créte 
et al. 1990) and identified through macroscopic and 
microscopic (10 — 25x) examination to family, genus 
or species whenever possible. Reference plant samples 
served for comparison during the identification process. 
Samples were grouped by taxon and their volume 
measured by water displacement (+0.25 mL). Plants 
were grouped into eight categories for statistical analy- 
ses: graminoids (Graminae and Cyperacae), farm crops, 
forbs (without flowers or fruits), leaves of shrubs and 
trees, wild flowers, wild fruits, mushrooms, and un- 
known. 

Forage quality — We reasoned that rumens of deer 
consuming poor quality forage would contain fewer 
nutrients with rapid assimilation and more nutrients 
with slow digestibility than counterparts feeding on 


362 


better quality forage. Poor quality forage could also 
include more secondary compounds slowing or in- 
hibiting digestion. We measured cell solubles, lignin, 
and phenolics as proxy for rapid digestibility, slow 
digestibility and defensive compounds. We also deter- 
mined nitrogen (N) content because it proved to be a 
good indicator of summer diet quality in Caribou 
(Rangifer tarandus) (Créte et al. 1990). We used the 
detergent method (Gauthier et al. 1989; Goering and 
Van Soest 1970) to measure cell solubles (1-NDF) and 
acid detergent lignin (ADL) (Mould and Robbins 
1982). Cell solubles are highly digestible whereas 
lignin is largely not (Robbins 1993: 251, 294). Total 
nitrogen content (N) was measured with an automat- 
ed Macro-Kjeldahl analyzer and total phenolic con- 
tent was determined by a calorimetric method using 
the Folin-Ciocalteu reagent (Marigo 1973; Sauvesry 
et al. 1991). Proteins are also highly digestible (Rob- 
bins 1993: 294) whereas phenolics includes 8000 sub- 
stances, of which tannins are known for reducing pro- 
tein digestion (Robbins 1993: 300). Some samples 
could not be analysed due to inadequate quantities. 
All analyses were duplicated or triplicated whenever 
a difference of more than 5% was found between the 
first two measurements. The average of the two clos- 
est estimates served as results. 

Forage availability — We estimated forage avail- 
ability during a companion study in the same study 
areas (Rouleau et al. 2002a). We measured forage bio- 
mass between the ground and 1.5 m in height at sites 
used by radio-collared deer in the two areas through- 
out the growing season with the help of 11 linear 
regression models. Vertical and lateral plant coverage 
over 2-m transects served as dependant variables in 
regression models to predict plant biomass by species 
(Rouleau et al. 2002b). 

Data analysis — In a first step, we determined 
whether sex (for deer >1 year old) and age (fawns vs. 
>1 year old) influenced forage composition or quality 
as summer progressed using MANOVA. We restricted 
the analysis to the HD area because our samples from 
deer with known sex and age were too small in the LD 
area. Analyses were performed on sex and age with 
respect to months, including sex by month, and age 
by month interactions. 

We used ANOVA for contrasting forage composi- 
tion and quality between the two study areas. We 
pooled months two by two for this analysis to ensure 
adequate sample size for each period in both regions. 
ANOVA were followed by pairwise protected LSD 
tests (PROC GLM, PDIFF option, SAS Institute Inc. 
1988). For all analyses, we ensured that residuals were 
normally distributed (Shapiro-Wilk test) and homog- 
enous (visual inspection of the plot). For forage com- 
position, we carried out ANOVA on ranks because we 
could not normalise residuals. We estimated means and 
their standard error with the LSMEANS statement 
(SAS Institute Inc. 1988). 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Results 

Rumen samples were collected from 93 deer (HD 
68 / LD 25): 64 adults, 11 fawns, and 18 of unknown 
age. On a volumetric basis, 80% of the particles 
examined could be identified to 68 food items. Males 
and females showed no overall differences in the food 
items they selected in the HD area when considering 
forage categories (F = 0.52; df = 6, 38; P = 0.79) or 
diet quality (F = 0.21; df = 5, 22; P = 0.95), without 
sex X age interaction for composition (F = 0.77; 
df = 24, 134; P = 0.76) and quality (F = 0.69; df = 20, 
74; P = 0.83). Neither did age influence forage com- 
position (F = 0.26; df = 6, 49; P = 0.95) or quality 
(F = 2.27; df = 5, 28; P = 0.08), nor did it interact with 
month on a composition (F = 2.31; df = 6, 53; P = 0.05) 
or quality (F = 1.43; df = 5, 28; P = 0.24) basis. Given 
these results, we did not take into account age and sex 
in subsequent analyses. 

Diet composition — Rumen contents differed between 
the two study areas (Figure 2). In spring, rumens con- 
tained no farm crops in LD but 13% in HD (P = 0.03). 
Grasses (P = 0.08) and flowers (P = 0.07) also tended 
to be more important food items in HD than in LD. 
Conversely, forbs (P = 0.04) and leaves of shrubs and 
trees (P = 0.12) accounted for a larger volume in LD 
samples. Diets during July-August were more-com- 
parable; the only significant difference was for forbs: 
32% in LD vs. 7% in HD (P = 0.01). Rumens collected 
during September and October contained more mush- 
rooms (P < 0.01) and leaves of shrubs and trees 
(P = 0.08) in LD, whereas those collected in HD includ- 
ed more fruits (P < 0.01) and grasses (P = 0.08). 

Differences between the two study areas were more 
striking when considering taxa for the whole growing 
season (Table 1). For this comparison, we eliminated 
fruits, mostly apples, because they were almost restrict- 
ed to HD and their more three-dimensional shape 
contrasting with that of most other two-dimensional 
plant tissues resulted in exceptionally large volume. 
Excluding fruits, 19 food items each accounted for 
more than 1% of the rumen volume in LD, summing 
up to 95 % of volume, compared with 20 food items 
making up 87 % of rumen content in HD. Forest forbs, 
such as Yellow Clintonia (Clintonia borealis) and Wild- 
Lily-of-the-Valley (Maianthemum canadense) were 
common food items in LD but rare in rumens collect- 
ed in HD. They were replaced in HD by forbs typical 
of openings and by farm crops, such as clover (77i- 
folium sp.) and alfalfa. Mountain Maple (Acer spica- 
tum) and Mountain Ash (Sorbus americana), the most 
common shrub leaves in rumens collected in LD, were 
replaced by Red Maple, Choke Cherry (Prunus vir- 
giniana), and Sugar Maple in HD; the latter species 
were also consumed by LD deer, but to a much lesser 
extent. Finally, apples, the most common item in HD 
rumens, were found in only one LD sample. 

Diet quality — Nitrogen (F = 0.91; df = 1, 52; P = 0.34) 
and phenolic content (F = 0.13; df = 1, 52; P = 0.71) 


2004 


DAIGLE, CRETE, LESAGE, OUELLET, AND Huot: SUMMER DIET OF DEER 363 


| Saint-Jean) 


| Lac 


70° 68° 66° 64° 
) Septlles aaa | 
| fe na 
| ae 
et nee 
lle “ieee, 


48° | 
| White-Tailed Deer 
rer! distribution Af 
|__| Study areas 


Ficure |. White-tailed Deer distribution in Québec and location of the two study areas, low density (LD) and high density (HD). 


did not differ between the two areas, with nitrogen 
remaining stable as the growing season progressed, 


| and phenolic content declining in September-October 


(Table 2). However, LD rumens contained more cell 
solubles (F = 5.66; df = 1, 69; P = 0.02) and lignin (F 
= 4.68; df = 1, 69; P = 0.03) than those from LD. 


Discussion 

Diet composition — Forage availability is reflected in 
diet composition. Although climate differed between 
LD and HD, the two regions shared most plant taxa 
and natural forage species had similar digestibility in 
both areas throughout the growing season (Lesage et 
al. 2000). However, biomass of preferred forage species 
found at sites used by deer was much larger in LD than 
HD (Rouleau et al. 2002a). Food items which domi- 
nated in LD diets remained almost absent from HD 
rumens. For instance Liliacae, such as Yellow Clintonia 
and Wild-Lily-of-the- Valley, which White-tailed Deer 
normally consume in the Northeast (Crawford 1982; 
Korschgen et al. 1980; McCaffery et al. 1974; Skinner 
and Telfer 1974; Waller and Alverson 1997) occupied 
a marginal volume in HD rumens. Liliacae are partic- 
ularly vulnerable to deer browsing (Créte et al. 2001; 
Waller and Alverson 1997) and can be used as indi- 


cators to estimate the impact of White-tailed Deer on 
plant communities (Balgooyen and Waller 1995). Wild- 
Lily-of-the-Valley was common in HD in the early 
1980s (Guérard and Legris 1984*). At the time of our 
study, its biomass was 20 times lower in HD than in 
LD (Rouleau et al. 2002a) and occurred as a marginal 
food item in the diet of HD deer (Table 1). Although 
not abundant on a volumetric basis, Wild-Lily-of-the- 
Valley occurred in 16% of the rumens collected in 
HD, showing that deer were still looking for this species 
even though its availability was largely reduced. Moun- 
tain Maple, which is known to be affected by deer 
(Balgooyen and Waller 1995), was a common species 
in HD in the past (Guérard and Legris 1984*), but 
seemed almost extirpated at the time of our study 
(Rouleau et al. 2002a). In LD, where natural forage is 
abundant (Rouleau et al. 2002a), Lesage et al. (2002) 
found that deer avoided cultivated fields. Our results 
support their findings and indicate that White-tailed 
Deer prefer to consume forest forages when they are 
readily available during the growing season. In HD, 
where most preferred food items had become scarce 
(Rouleau et al. 2002a; Boucher 2004), deer switched 
to other natural forage still persisting (e.g., Red Maple, 
Choke Cherry and Sugar Maple) and relied on farm 


364 THE CANADIAN FIELD-NATURALIST Vol. 118 


May-June 


GLD 
gm HD 


Farm crops Forbs Flowers Fruits Grasses Leaves Mushrooms 


July-August 


Farm crops Forbs Flowers Fruits Grasses Leaves Mushrooms 


September-October 


Farm crops Forbs Flowers Fruits Grasses Leaves Mushrooms 


FIGURE 2. Composition (% volume + SE) of rumen samples collected in a low-density LD (n = 25) and a high-density HD 
(n = 68) area of southern Québec during the growing seasons of 1997 and 1998. 


crops for a large part of their summer diet. Comparison Fruits, mostly apples (90%), were a major compo- 
of forage availability and consumption (Table 1) sug- __ nent of the fall diet in HD. Our results support the idea 
gests that Red Maple, Choke Cherry, and Sugar Maple that fruits are important food items where and when 
suffered high browsing pressure in HD. they are available (McCaffery et al. 1974; Short 1971; 


2004 


DAIGLE, CRETE, LESAGE, OUELLET, AND HUoT: SUMMER DIET OF DEER 365 


| TABLE |. Most common food items (volume >1%) found in White-tailed Deer rumens from a low density LD (n=23) and a 


high density HD area (n=63) of southern Québec during the growing seasons of 1997 and 1998. 


Skinner and Telfer 1974). Apple trees and oak are 
absent from forests in LD. We assumed that apples 
found in one LD sample came from baits set by hunters. 
Baiting was also common in HD, but we think that 
apples ingested by deer in this region primarily origi- 
nated from uncultivated apple trees growing in young 
forests based on the fact that most specimens found 
in rumens were smaller than commercial apples. 
Mushrooms were a common food item in both study 
areas. Like apples, but to a lesser extent, their shape and 
size may have exaggerated their relative importance 
on a volumetric basis. Their frequent occurrence in 
rumens indicates that they represented a common food 
item for White-tailed Deer. Mushrooms were consumed 
mostly in spring in HD and in fall in LD, which may 
be explained by different seasonal availability or by the 
possibility that White-tailed Deer feed on them when 
preferred foods are rare at both ends of the growing 
season. Many studies reported large amounts of mush- 


| % volume % occurence Availability (g/m) 
| Food items Category LD HD LD HD LD HD 
Leaves, stems, and grain 
| Acer spicatum Leaf 26.9 0.6 30.4 4.8 19.0 <0.) 
Fungi Mushroom 14.7 16.4 26.1 20.6 <0. 1 <0.) 
| Botrychium sp. Forb a2, 0 4.3 0 
| Clintonia borealis Forb 6.2 0.1 34.8 1.6 Dei 0.1 
| Maianthemum canadense Forb 5.9 Ie) 47.8 15.9 el 0.2 
| Sorbus americana Leat 4.4 0 26.1 0 <0.1 
| Fraxinus nigra Leaf 4.0 0 4.3 0 
| Epilobium angustifolium Forb 3.6 0 PAVE 0 0.8 
Prunus virginiana Leaf 3.6 8.9 Paved 28.6 <0.1 <0.1 
Prunus pennsylvanica Leaf pos) 0.9 26.1 4.8 <0.1 
| Trillium sp. Forb PRS} 0.2 13.0 6.3 <0.1 <0.1 
Aralia nudicaulis Forb 23 2.1 8.7 6.3 2.3 <0.1 
» Acer rubrum Leaf 1.9 10.3 PSH 41.3 2.6 0.1 
| Taraxacum officinale Forb 19 2D 13.0 Vas7) 0.9 0.2 
Populus tremuloides Leaf od Ie 13.0 95 4.3 0.1 
| Graminae Grass ites) 8.4 43.5 76.2 2.0 
| Acer saccharum Leaf 13 6.6 13.0 14.3 1.6 <0.1 
Taxus canadensis Leaf 1.3 <0.1 4.3 1.6 
| Salix sp. Leaf Les 0.5 8.7 4.8 <0.1 
| Solidago sp. Forb 0 1.2 0 4.8 
| Fragaria sp. Forb 0 1.5 0 19.0 0.1 
_ Lotus corniculatus Farm crop 0 1.6 0 1.6 
| Prunus serotina Leaf 0 1) 0 Hilal 0.3 
| Sonchus sp. Forb 0.2 1.9 4.3 322 <0.1 
| Rubus sp. Leaf 0.7 23 8.7 27.0 
| Potemogeton sp. Forb 0 3.0 0 32 
| Malus pumila Leaf 0 32 0 6.3 
| Zea mays Farm crop 0 3.6 0 7.9 
Medicago sativa Farm crop 0 3.8 0 7.9 
Trifolium sp. Farm crop 0.1 4.8 4.3 20.6 
Fruits 
Malus pumila Fruit 100 90.1 4.3 36.5 <0.1 <0.1 
Quercus sp. Fruit 0 3.6 0 1.6 
Fragaria sp. Farm crop 0 Sal 0 1.6 
Crataegus sp. Fruit 0 22 0 6.3 


rooms in White-tailed Deer diet (Crawford 1982; John- 
son et al. 1995; Korschgen et al. 1980; McCaffery et 
al. 1974), but nobody has yet considered the potential 
impact of deer herbivory on mushrooms. 

Diet quality - LD deer had access to a greater bio- 
mass of preferred forage (Rouleau et al. 2002a) and 
consumed more forbs and leaves of shrubs and trees 
than HD deer, which compensated by ingesting farm 
crops, grasses and fruits. Different forage intake result- 
ed in a diet containing more cell solubles and lignin 
in LD than in HD deer. Rumen contents did not differ 
with respect to phenolics, which may have reflected 
the complex nature of the chemical group (Robbins 
1993: 253). Protein content followed a similar trend 
as that for cell solubles, although the difference between 
LD and HD was small and non-significant. Between 
May and September, cell solubles were 8% greater in 
LD than HD rumens. Cell solubles, which are composed 
of sugars, protein, non-protein nitrogen, lipid, organic 


366 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE 2. Mean (SE; n) content (%) of cell solubles, lignin, phenolics, and nitrogen in White-tailed Deer rumen samples collected 
in a high-density (HD) and a low-density (LD) study area of southern Québec during the growing seasons of 1997 and 1998. 


May-June 

HD LD 

Cell solubles 32.4 40.7 
(1.4; 21) (2.0; 14) 

Lignin 222 23.3 
(1.0; 21) (1.2; 14) 

Phenolics 3i5) 3.6 
(0.1; 16) (0.2; 13) 

Nitrogen Te 7.0 
(0.3; 16) (0.3; 13) 


acid and soluble minerals (Iason and Van Wieren 1999), 
are highly and rapidly digestible, whereas cell walls (1- 
cell solubles) are only partly digestible and necessitate 
some retention in the rumen for microbes to degrade 
them (Iason and Van Wieren 1999). LD does benefit- 
ed therefore from a richer diet than HD counterparts 
during the end of gestation and peak lactation, a peri- 
od with elevated energy demand in cervids (Mauget 
et al. 1997). In late summer, the high consumption of 
apples, which contain little fiber (Robbins 1993), can 
explain the improved quality of rumen contents in HD. 
The higher concentration of lignin in LD than in HD 
rumens likely came from the greater consumption of 
farm crops and grasses in the latter area, which have 
a lower level of lignin than natural forages do (Rob- 
bins et al. 1987). 

The difference in forage quality likely reduced the 
growth and condition of White-tailed Deer in HD, 
where strong competition for forage has prevailed. 
Deer density in HD increased by 10 fold since the late 
1970s, whereas it remained relatively low and stable 
in LD (Lesage et al. 2001). Deer had the same size in 
both study areas before 1987 (Potvin 1989), but their 
body mass progressively decreased in HD (Lesage et 
al. 2001; Potvin 1994*). At the time of our study, the 
eviscerated carcass mass of fully grown male deer aver- 
aged 80 vs. 116 kg in HD and LD, respectively (Lesage 
et al. 2001). Additional measurements throughout the 
deer range in Québec revealed that deer size began to 
decline when forage competition intensified in sum- 
mer; 1.e., forage availability <10 000 kg/deer (Boucher 
2004). Our results concur with the hypothesis that rural 
deer living at high density cannot fully compensate for 
the rarity of natural forage in woodlots during summer 
by consuming cultivated plants (Rouleau et al 2002a). 
All of these studies suggest that deer populations have 
reached excessive densities in HD. 


Acknowledgments 

We thank R. Bélanger, J. Boisvert, B. Deschamps, 
and M. Desrochers from Transports Québec, who kind- 
ly gave us access to deer carcasses. J. Bachand, C. 
Bélanger, M. Bélanger, N. Bergeron and E. Jeanson 
helped in collecting the rumen samples. We are grateful 
to G. Picard for conducting the chemical analysis, to 


July-August September-October 


HD LD HD LD 
34.1 42.7 40.7 3919 
(1.8; 11) (2.1; 4) @i5223) @a72) 
21.9 23.4 17.8 24.0 
(1.4; 11) (2.3; 4) (1.0; 23) (6322) 
3.4 Bi) 2.4 Di 
(0.2; 7) (0.3; 4) (0.1; 16) (0.4; 2) 
6.8 6.5 6.9 7.0 
(0.4; 7) (0.6; 4) (0.3; 16) (0.8; 2) 


N. Desrosiers, and E. Normand for their assistance in 
the laboratory, and to C. Roy for helping with plant 
identification. G. Daigle, from the Service de consul- 
tation statistique, Université Laval, did the statistical 
analysis. 


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Received 31 March 2003 
Accepted 1 November 2004 


Winter Habitat Use by Wolves, Canis lupus, in Relation to Forest 
Harvesting in West-central Alberta 


GERALD W. Kuzyk!, JEFF KNETEMAN?, and FIONA K. A. SCHMIEGELOW! 


' Department of Renewable Resources, University of Alberta, Edmonton, Alberta, T9H 4N1 Canada 
> Fish and Wildlife Division, Alberta Sustainable Resource Development, Hinton, Alberta, T7V 2E6 Canada 
> Corresponding author: gkuzyk @ualberta.ca 


Kuzyk, Gerald W., Jeff Kneteman, and Fiona K.A. Schmiegelow. 2004. Winter habitat use by Wolves, Canis lupus, in relation 
to forest harvesting in west-central Alberta. Canadian Field-Naturalist 118(3): 368-375. 


Forested landscapes in west-central Alberta are facing increased pressures from forest harvesting and other land-use 
activities, which may alter the movements and distribution of Wolves and ungulates. Information on habitat use by Wolves in 
logged forests is scarce, potentially limiting effective land-use planning in the boreal forest. Nine Wolves, from four Wolf 
packs, were fitted with GPS radiocollars in the Rocky Mountain foothills, near Grande Cache, Alberta (2000-2001). We 
found Wolves did not use the landscape randomly, but rather exhibited a significant preference for non-forested natural 
habitats (shrubs, water), relative to their availability. Within forest habitats, Wolves used cutblocks proportionately more than 
unharvested forest and non-forested anthropogenic habitats (pipelines, clearings); however, selection of forest cutblocks was 
not statistically significant. We found no evidence that Wolves preferred or avoided forest cutblock edges. Wolf pack 
territories contained various levels of timber harvesting, but most areas were still in the early stages of harvest. Nevertheless, 
these areas have been allocated for large-scale harvesting. Understanding the potential responses of Wolves to rapidly 
changing landscape mosaics poses a significant challenge to researchers and managers, but such information is important to 


informing future land-management and conservation strategies for boreal forest Wolf-prey systems. 


Key Words: Wolf, Canis lupus, Caribou, Rangifer tarandus, forestry, habitat, Moose, Alces alces, predation, Alberta. 


Much of the world’s boreal forest is undergoing in- 
creased demands from resource extraction industries, 
where related activities such as forest harvesting may 
alter habitat use by large carnivores (McLellan and 
Hovey 2001; White et al. 2001). Wolves (Canis lupus) 
are a common predator of ungulates in Canada’s for- 
ests, yet there are few data on how their use of habitat 
might be affected by forest harvesting (Jedrzejewska 
et al. 1994; Kohira and Rexstad 1997; Kunkel and 
Pletscher 2000). Logging of forests can change the spa- 
tial dynamics of Wolves and their prey, resulting in 
conflict between resource development and wildlife 
management (Hervieux et al. 1996). For example, in 
Alberta, Woodland Caribou (Rangifer tarandus caribou) 
are classed as a threatened species (Edmonds 1998), 
and Wolf predation is considered a primary reason for 
their decline (Edmonds 1988; McLoughlin et al. 2003). 
It is therefore important to understand how Wolves 
respond to forest harvesting, as habitat use by Wolves 
may change in response to logging activities, and could 
affect predation risk to Caribou and other ungulates. 

Forest harvesting can cause habitat fragmentation 
and alter predator-prey systems. Predators may follow 
habitat edges due to ease of travel (Bider 1968). As 
well, when patch size decreases, predator numbers may 
increase due to increased prey density and diversity 
(Gates and Gysel 1978; Yahner 1988). For example, 
Red Fox (Vulpes vulpes) and Coyote (Canis latrans) 
densities can increase with more landscape fragmenta- 
tion, and habitat edges are favored for hunting (Oehler 
and Litvaitis 1996). 


In a Wolf-prey system in Canada’s boreal forest, 
forestry activities have the potential to alter the preda- 
tion risk to ungulates from Wolves in three ways. First, 
differential spatial and temporal habitat selection sep- 
arates Woodland Caribou and Moose (Alces alces) 
distribution, reducing the risk of predation to Caribou 
from Wolves (Bergerud and Elliot 1986). Caribou sel- 
ection for higher elevations reduces the chance of en- 
counter by Wolves hunting Moose, a primary prey spe- 
cies, in lower elevations riparian areas (Seip 1992). If 
the spatial separation of Caribou and Moose is altered 
by logging roads and forest cutblocks, it has been ar- 
gued that Wolves will have increased access to, and 
greater encounter rates with Caribou, resulting in a Cari- 
bou decline (Bergerud 1988; Seip 1992). In northeast 
Alberta, linear developments (roads, seismic lines, 
trails) were found to affect the spatial separation be- 
tween Caribou and Moose, where linear corridors en- 
hanced Wolf travel efficiency (James 1999) and Cari- 
bou mortalities caused by Wolves were found closer 
to linear corridors than expected by chance (James 
and Stuart-Smith 2000). 

Second, Caribou select for older forests (Szkorupa 
2002) and distribute at low densities (Bergerud 1988). 
Caribou spatial overlap with Wolves is reduced and, 
correspondingly, so too is the risk of detection and pre- 
dation. Reducing the amount and patch size of older 
forest by timber harvest may temporarily increase 
Caribou densities (Bergerud 1988), and Moose, deer 
(Odocoileus spp.) and Elk (Cervus elaphus) may use 
these remaining patches for cover. Higher densities of 


368 


2004 


Caribou in more constricted areas, and wider distribu- 
tion of Wolves in response to primary prey distribution, 
may increase the chances of Caribou being detected 
by Wolves. 

Third, Moose, Elk and deer are attracted to recently 
logged areas that support high quality regenerating 
forage (Peek et al. 1976; Tomm et al. 1981; Stelfox et 
al. 2001). This represents a concentrated prey base for 
Wolves, which may influence how Wolves use land- 
scapes. If Wolves frequent forest cutblocks searching 
for Moose, Elk and deer, and if cutblocks occur near 
preferred Caribou habitats, this may increase predation 
risk to Caribou. All three outcomes potentially affect- 
ing Caribou depend on information about Wolf behav- 
ior and use of changing landscapes. 

Forest harvesting in west-central Alberta was large- 
ly initiated in the late 1960s and has accelerated in 
recent years. Energy sector activities (oil and gas ex- 
ploration and development) are also altering these 
landscapes, resulting in cumulative land use impacts 
(Hervieux et al. 1996). For decisions concerning long 
term wildlife conservation, resource managers and 
land-use planners require new information about how 
Wolves use habitat in logged forests and under chang- 
ing landscape conditions. 


ALBERTA 


Study area 
shown indetai 


400 Kilomaers 
7 


gs Wolf locations (GPS) 
© Wolf pack territory 


». Town of Grande Cache 
— Major Rivers 


Zones 


Data Source: Alberta Environment Natural Resources, 
Govemment of Canada 2000; DEM published with permission of 
Alberta Sustainable Resource Development. Natural Resource 
Canada and the Spatial Data Warehouse. Weyerhaeuser Company. 


KUZYK, KNETEMAN, and SCHMIEGELOW: HABITAT USE BY WOLVES 


369 


We used Global Positioning Systems (GPS) radio- 
collar technology to examine winter habitat use of 
Wolves in west-central Alberta. We chose to examine 
fine-scale Wolf movements that correspond with John- 
son’s (1980) third order habitat selection: movements 
of animals within their home range. We examined two 
questions. First, do Wolves use forest cutblocks prefer- 
entially over other habitat types? Second, do Wolves 
prefer cutblock edges? We predicted that Wolves would 
prefer forest cutblocks over other habitats, due to the 
expected increase in ungulate densities in regenerating 
forests. We also predicted Wolves would prefer forest 
cutblock edges relative to areas farther away from them, 
due to ungulate use of cutblock edges for feeding and 
proximity to cover (Stelfox et al. 2001*). 


Study Area 

The study area is approximately 5000 square kilo- 
meters, located in the foothills of west-central Alberta, 
near the town of Grande Cache (54°N 119'W) (Fig- 
ure 1). The area is classed into subalpine and boreal 
natural subregions (Beckingham and Archibald 1996), 
and contains several main rivers and a dendritic pat- 
tern of creeks; lakes are scarce. Elevations range from 
1300-1800 meters, and the climate is subarctic, with 


Map compied by S. Shirkoffand G. Kuzyk. Unwersty of Alberta 


FiGure 1. Distribution of four Wolf packs monitored in winters of 2000 and 2001 in west-central Alberta. 


370 


short wet summers and long cold winters. Temperatures 
average 16°C in July and -13.5°C in December (Beck- 
ingham and Archibald 1996). The forests are primarily 
Lodgepole Pine (Pinus contorta) and some White 
Spruce (Picea glauca). The wetland complexes support 
mostly Black Spruce (Picea mariana) and some Tam- 
arack (Larix laricina). Some south facing slopes sup- 
port Trembling Aspen (Populus tremuloides) and wil- 
low (Salix spp.). 

This area supports a high diversity of large mam- 
mals: Woodland Caribou, Moose, Elk, Mule Deer 
(Odocoileus hemionus), White-tailed Deer (Odocoileus 
virginianus), Bighorn Sheep (Ovis canadensis), Moun- 
tain Goats (Oreamnos americanus) and Wild Horses 
(Equus cabalus). Wolves, Coyotes (Canis latrans), 
Grizzly Bears (Ursus arctos), Black Bears (Ursus 
americanus) and Cougars (Felis concolor) also exist 
throughout the study area. 

Major land use activities include forest harvesting, 
oil and gas exploration and development, coal mining, 
commercial trapping, and public uses such as hunting, 
fishing, hiking, horse packing and camping. Access 
is primarily on roads created for resource extraction, 
pipelines and seismic lines. Further descriptions of 
the study area can be found in Edmonds (1988) and 
Smith et al. (2000). 


Wolf Location Data 

Nine Wolves in four packs were captured and fitted 
with GPS radiocollars in winters of 2000 and 2001 
(Table 1). Three packs were located in areas with a 
migratory mountain Caribou population and one pack 
with a sedentary boreal Caribou population. All Wolf 
handling was approved by the Faculty of Agriculture, 
Forestry and Home Economics Animal Care Policy 
(Number 96-99D), subject to the protocols of the 
Canadian Council of Animal Welfare. Wolf captures 
were accomplished by helicopter darting (Ballard et al. 
1991) or netgunning, then physically restraining the 
Wolf with restraining forks, and hand-injecting 1-2 mls 
of telazol at 200 mg/ml (Kuzyk 2002a). Wolves were 
fitted with store aboard GPS radiocollars (Lotek En- 
gineering Sytems, Newmarket, Ontario). In the winter 
of 2001, the Prairie Creek and Cutbank packs each had 
two members with GPS radiocollars. To avoid pseudo- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


replication (Hurlburt 1984), location data and asso- 
ciated patterns of habitat use from these individuals 
were averaged for their respective packs (Table 1). 

As this study was designed to understand Wolf hab- 
itat use in winter, the following criteria were used to 
select Wolf location data: 

(1) Location data from 31 January to 25 April 25 in 2000 
and 2001 were used for analyses. These dates were used for 
two reasons: first, most Caribou in the study area (the migra- 
tory mountain ecotype) leave the forested foothills in late 
winter and spring to calve in the nearby mountains (Edmonds 
1988); second, a spring cutoff time also has ecological rele- 
vance to Wolves. In spring near whelping time, Wolves are 
thought to change their hunting patterns by switching from 
hunting as a pack and preying on ungulates, to hunting 
alone or in small units in search of smaller prey, with their 
activities centering on the den and pups (Mech 1970). There- 
fore a single GPS collared Wolf would no longer represent 
the behavior of their pack, and would not meet our design 
criteria. 

(2) To provide consistency in GPS collar programming, 
six-hour locations were chosen (4 per day) as the minimum 
common sampling unit for analysis. Wolves are sporadic in 
their movements, and may travel at rates of about 8 km/hr 
while hunting (Mech 1966), or relatively short distances when 
near a killsite (Kuzyk 2002a). When near a killsite, they sel- 
dom rest in one location for periods longer than six hours 
(Mech 1970). 

(3) Wolf locations outside calculated pack territories were 
not used. These Wolves were assumed to be dispersing from 
their natal territory and thus behaving differently from their 
pack (Gese and Mech 1991). 

Data were differentially corrected using N4Win 
Version 2.40 program (Lotek Engineering Inc. 2000) 
and were assumed accurate within 14 meters, 95% of 
the time (Lotek Engineering Inc. 2000). Wolf locations 
with Dilution of Precision (DOP) values greater than 
15 were removed from the analysis (<2 % of total loca- 
tions). High DOP values and radiocollar malfunctions 
made for unequal locations per Wolf pack (range 152 
to 279) over the duration of this study (Table 1). 


Habitat Classification and GIS Methods 
Wolf location data were imported into ArcView 
Version 3.1 (Environmental Systems Research Insti- 
tute Inc., Redlands, California). Current, digital forest 
inventory coverages were obtained from Weyerhauser 
Canada Limited, Canadian Forest-Products and Al- 


TABLE |. Wolf packs with associated number of GPS locations and area of habitat use (territory size) in west-central Alberta 


in winters of 2000 and 2001. 


Wolf Pack Wolf Year Dates Number of Locations Area (km?) 
Cutbank W5 2000 24 January — 16 March 185 714 
Prairie Creek W9 2000 28 January — 25 April 252 286 
Simonette W13 2000 31 January — 25 April 279 786 
*Prairie Creek W5 and W9 2001 18 February — 25 April 231 182 
*Cutbank W19andW21 = 2001 15 February — 25 April 258 448 
Lynx Creek W22 2001 15 February — 25 April 247 1848 
Simonette W30 2001 17 February — 4 April 152 398 


* Locations were averaged for two collared Wolves which belonged to the same pack. 


2004 


KUZYK, KNETEMAN, and SCHMIEGELOW: HABITAT USE BY WOLVES 


37] 


TABLE 2. Description of habitat categories used in compositional analysis for Wolf packs in west-central Alberta during late 


winters of 2000 and 2001. 


Habitat 2 


Unharvested forest 
all harvestable forest 
burn 


Habitat | 


Forest cutblocks 
all harvested forest 


Habitat 3 Habitat 4 

Non-forest (natural) Non-forest (anthropogenic) 
herbaceous grassland clearing 

sand, flooded land right-of-way 

closed and open shrub industrial infrastructure 
coniferous scrub pipelines 


deciduous scrub 
brush, windfall 

open and treed muskeg 
water 


geophysical 
perennial forest crops* 


*Perennial forest crops are denoted as anthropogenic by the forest companies and account for < 0.6km?* of one Wolf pack’s 


territory. (Simonette pack — total territory size is 786 km7). 


TABLE 3. The percentage of use (GPS locations) and availability (area in km?) of five habitat categories for Wolf pack 
territories in west-central Alberta during late winters of 2000 and 2001. 


Cutblock Forest Non-forest* Non-forest* Non-forest 

Shrubs Water Anthropogenic 

Wolf Pack Year Used_1 Avail_1 Used_2 Avail_2 Used_3 Avail_3 Used_4 Avail_4 Used_5 Avail_5 
Cutbank 2000 34.1 28.9 57.8 64.6 7.0 A: 1.1 0.3 0.0 0.8 
Prairie Creek 2000 Lieve Aes) 77.8 80.2 6.7 4.8 LED, 1.0 2.4 Hes) 
Simonette 2000 PN SYS) 61.6 78.8 Gs 3.0 DES 0.6 TD 1.8 
Prairie Creek 2001 6.3 5.9 81.2 87.1 9.1 4.1 fe 1.1 M7 1.8 
Cutbank 2001 Asay SD) 35/9 60.0 20.2 3.4 0.4 0.3 0.4 0.8 
Lynx Creek 2001 0.8 5.4 1S} 85.9 202 8.5 0.8 0.1 0.8 0.2 
Simonette 2001 16:4: - 1420 68.4 78.3 113) 5) 0.7 0.5 1.3 Le, 
Total Mean 19.1 16.9 65.7 76.4 1220 5.0 le, 0.6 2.0 1D, 
SE S27, 4.3 6.0 3.9 23, 0.7 0.3 0.1 0.9 0.2 


*Non-forest natural is subdivided into shrub and water categories for descriptive purposes only. 


berta Government Phase 3. The minimum mapping 
unit for these coverages was | ha for forest polygons. 
However, the resolution of line features, such as roads, 
was much greater, as these were spatially referenced 
from high-resolution remotely sensed data (~5 m res- 
olution), or from GPS readings taken in the field. Thus, 
the resolution of our animal location data was com- 
mensurate with the resolution of the landscape cover- 
ages used, for purposes of evaluating coarse-scale habi- 
tat use. Minimum convex polygons (MCPs) of Wolf 
territories were initially calculated with an animal 
movement extension in ArcView (Hooge and Eichen- 
laub 1997). MCPs were considered an appropriate 
method for delineating general territory boundaries 
in order to evaluate coarse-scale habitat use. Due to a 
small portion of the GIS coverages missing within each 
Wolf pack territory, territory sizes for subsequent 
analyses were adjusted by summing all the areas within 
the MCPs for which we had GIS coverages (Table 1). 

As Wolves live in a defined territory (Mech 1970), 
each territory was classified into four habitat categories 
to reflect coarse scale patterns of use. These categories 
were: (1) forest cutblocks, (2) unharvested forest, (3) 
non-forest natural (shrubs and water) and (4) non- 


forest anthropogenic (pipelines, wellsites) (Table 2). 
The area of non-forest natural was divided into “shrub” 
and “water” classes for descriptive purposes (Table 
3), but the data were pooled for analysis. As the focus 
of this study was to determine habitat use of Wolves 
in managed landscapes, a further analysis was con- 
ducted to determine Wolf use of forest cutblock edges. 
Forest cutblocks were buffered using specified dis- 
tances starting from the edge of the forest-cutblock and 
proceeding into the forest. We did not use locations 
inside cutblocks in this analysis. Buffer distances were 
consistent with those studying Caribou avoidance of 
linear features (Dyer et al. 2001; Oberg 2001), starting 
from the edge of the cutblock to 100 m, 101-250 m, 
251-500 m, 501-1000 m and >1000 m. The category 
of >1000 m was also chosen as the farthest distance 
for comparison to Smith et al. (2000), who found that 
Caribou in west-central Alberta may avoid cutblocks 
by about 1200 m. 


Statistical Analysis 

Compositional analysis (Aebischer et al. 1993) was 
conducted by integrating Wolf GPS location data and 
forest inventory data within a GIS (ArcView 3.1) to 


B72 


determine if there was a preference in Wolf use of habi- 
tat or buffer categories. Aebischer et al. (1993) suggest 
a minimum of six radiotagged animals are required to 
perform compositional analysis, and replication across 
years is acceptable. Therefore, our sample of seven 
Wolves over two winters was adequate for this test. 
Compositional analysis compares the amount of “used 
habitat” to the amount of “available habitat” and tests 
whether habitats are preferred or avoided more than 
expected by random (Johnson 1980). The number of 
Wolf locations in each habitat or buffer category rep- 
resented used habitat. The available habitat was the 
total area of each habitat or buffer category (Table 3). 
If there was no use of a habitat category, 0% use was 
replaced with 0.001%, as this represented a value lower 
than the smallest recorded nonzero percentage (Aebis- 
cher et al. 1993). 

A chi-square test was used to determine if Wolf use 
of habitat or buffer categories was significantly non- 
random, then each habitat category was ranked in terms 
of its use. To determine if any habitats were selected 
over others, a difference for each pair-wise comparison 
was calculated using log ratios. This compared each 
habitat category within each Wolf pack territory. The 
means and standard errors for each comparison were 
calculated across all Wolf packs, and the pair-wise 
differences were tested for significance using a t-test 
(Aebischer et al. 1993). Alpha level for all tests was 
set at 0.05. 


Results 
Wolf Habitat Use 

Territory size for the four Wolf packs ranged from 
182 — 1848 km? (Table 1). The availability of each of 
the four habitat categories varied markedly: the per- 
centage of forest averaged 76.40% (SE 3.90) for all 
packs, and thus was the most dominant habitat, where- 
as non-forest anthropogenic cover averaged only 1.23% 
(SE 0.25) of available habitat across Wolf territories 
(Table 3). Wolves showed a significant deviation from 
random use of the four habitat types (v7 = 7.815, df = 3, 
p = 0.036) selecting non-forest natural (shrubs-water) 
habitats over both forest (t = -4.281, df = 6, p = 0.005) 
and cutblocks (t = -2.92, df = 6, p = 0.027), in relation 
to their availability (Table 4). No other pair-wise com- 
parisons were significant. However, when ranked in 
preference by habitat type, forest cutblocks were pref- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


erred more than both forest and non-forest anthropo- 
genic areas. 


Wolf Response to Forest Cutblock Edges 

The available areas for all distance buffers less than 
1000 m were similar, with variation due mostly to 
dissolving buffers for adjacent cutblocks. Wolf use of 
distance buffers did not deviate significantly from ran- 
dom (x7 = 2.349, df = 3, p = 0.503). When buffer dis- 
tances were compared using compositional analysis, 
no significant difference was found between distance 
categories related to forest cutblock edges. When 
ranked, the 501-1000 m buffer distance was the most 
preferred, followed by the 0-100m buffer, with the 
least preferred being the buffer >1000 m. 


Discussion 

Wolves have been described as habitat generalists 
(Mech 1970; Mladenoff et al. 1995). On a coarse spa- 
tial scale, Wolves inhabit large tracts of forest (Mech 
1995) and may prefer mixed wood forests over either 
homogenous coniferous or deciduous forests (Mladen- 
off et al. 1995; Krizan 1997). Wolves may use forests 
altered by logging, as these areas provide good deer 
habitat, and thus support an important prey base for 
Wolves (Mladenoff and Sickley 1998). In this study, 
GPS radiocollar technology allowed for a more re- 
fined examination of Wolf habitat preferences, show- 
ing Wolves in our study area do not use the landscape 
randomly. In general, Wolves preferred habitats with 
young vegetation, in both non-forest natural habitats 
and forest cutblocks. This is consistent with increased 
ungulate abundance in areas of young vegetation (Peek 
et al. 1976; Stelfox et al. 2001*), which attract Wolves 
(Bergerud 1988). However, increased road access into 
these areas may also allow humans to alter activity 
patterns of Wolves (Theuerkauf et al. 2003), or affect 
Wolf numbers by direct or indirect killing (Mech 
1995, Mladenoff and Sickley 1998). In this study, the 
least used habitat by Wolves was non-forest anthro- 
pogenic (pipelines, right-of-ways), possibly to avoid 
human contact. Two radio-collared Wolves were known 
to have been shot during this study, and several other 
collars were lost to unknown factors (Kuzyk 2002a). 

We found little support for our first prediction that 
Wolves select forest cutblocks. Wolves did use cut- 
blocks proportionately more than their availability and 
were ranked above forest or anthropogenic features, 


TABLE 4. Results from compositional analysis (p values in parenthesis; + denotes row > column and — column > row) for 
comparing four habitat categories for four Wolf packs in west-central Alberta during late winters of 2000 and 2001. 


1 2) 
Cutblock Forest 
1 Cutblock +(0.947) 
2 Forest - 
3 Non-for_natural + + 
4 Non-for_anthro. - - 


* denotes significance at (p<0.05). 


3 4 
Non-for_natural Non-for_anthropogenic 
-(0.027) * +(0.902) 
-(0.005) * +(0.903) 
+(0.177) 


— 2004 


| but differences were not significant. Similarly, Wolves 
_ in Ontario were found to use cutblocks in proportion 
to their occurrence (Krizan 1997). We had the advan- 


tage of GPS radiocollars which allowed for a large 
collection of location data, compared with the traditi- 
onal VHE collars used by Krizan (1997). However, we 
acknowledge that our analyses still lacked statistical 
power, due to the relatively small sample of Wolves 
radiocollared. 

The amount of harvested forest differed substan- 


| tially between Wolf packs and may have accounted 


| 


for variation in Wolf use of forest cutblocks. The Lynx 
Creek pack had only 5% of its territory as forest cut- 
blocks, whereas 36% of the Cutbank pack’s territory 
consisted of recent cutblocks. This seven-fold differ- 
ence in the amount of harvested forest between packs 
may have influenced habitat preferences. Kohira and 
Rexstad (1997) found no evidence that Wolf diets 
differed between logged and unlogged areas in the 
coastal rainforests of Alaska. About 6% of that total 
study area was logged, with the amount of area logged 
ranging from 1-26% between Wolf pack territories. 
This differs from our study area, where about 17%, 
or approximately three times as much area has been 
logged. In southeast British Columbia, researchers also 
did not find evidence that forest harvesting increased 
the vulnerability of Moose to predation by Wolves, 
where about 13% of the area was logged (Kunkel and 
Pletscher 2000). 

Wolves in our study did show a significant prefer- 
ence for non-forested natural habitats (shrubs/water) 
over cutblocks and forest, which might be explained 
by a number of selection criteria. Wolves prefer to 
rest in open areas, and may travel several kilometers 
to reach such preferred sites (Mech 1970). Wolves in 
this study area were observed on numerous occasions 
resting in open meadows, muskegs, hillsides and beav- 
er ponds, often when they were near killsites (Kuzyk 
2002a). The shrubs in these habitats have open crowns, 
which allows both penetration of sunlight and structure 
for protection from the wind, thus providing Wolves 
cover while resting. 

The shrubs in this non-forest habitat type also pro- 
vide forage and cover for ungulates. During this study, 
Wolves made deer kills in shrubby willow areas, and 
Moose, deer and Elk kills in or near riparian areas 
(Kuzyk 2002a). Elk are primarily grazers, and may be 
attracted to these shrub patches due to the increased 
availability of grasses. Bjorge and Gunson (1989), in a 
nearby Wolf study, noted that Elk, especially Elk calves, 
were a preferred prey for Wolves in winter. During the 
limited kill rate work associated with this study (Kuz- 
yk 2002a), only one cow Elk kill was documented. In 
Jasper National Park, Wolves hunt deer while mov- 
ing to pockets of Elk (Carbyn 1974; Weaver 1994). It 
is possible that shrubby areas do represent reliable 
patches of prey, and the Wolves investigate them for 
prey regularly. 


KUZYK, KNETEMAN, and SCHMIEGELOW: HABITAT USE BY WOLVES 


373, 


Water was also included in this preferred habitat 
class. It is common for Wolves to use frozen waterways 
as travel routes where snow is most compacted and 
ice makes travel easy. In winter Wolves probably use 
creeks and rivers to travel among ungulate winter 
ranges. Also there is an increased chance of encounter- 
ing Moose that use riparian areas in winter (Hayes et 
al. 2000) and Wolves are known to frequently kill 
ungulates on iced surfaces (Mech 1991). 

Our second prediction of Wolf preference for for- 
est cutblock edges was also not supported. There was 
no significant difference in Wolf preference for any 
buffer distance categories, nor was the 0-100 m buffer 
class ranked highest. Habitat was not controlled for 
in the buffer categories, which may have confounded 
the analysis. The behaviors of Wolves, such as feeding 
at killsites, resting and hunting may also be diluting 
the effect of any preference or avoidance of cutblock 
edges. Wolves hunt a diversity of prey, and chase dis- 
tance varies with each prey type (Paquet 1989). For 
example, the average chase distance for a Moose is 
883 m (Paquet 1989). Depending on where Wolves lo- 
cate them, Moose could choose to run to the nearest 
forest to avoid attacking Wolves (Stephens and Peter- 
son 1984), or remain stationary and aggressive (Mech 
1966), or stationary and non-aggressive (Kuzyk 2002b), 
and still avoid attack by Wolves. These results, and 
those related to broader habitat selection questions, 
suggest that consideration of behaviors associated with 
different habitat types is an important component of 
interpretation in Wolf habitat use studies. 

Forest harvesting alters both the amount and spatial 
distribution of habitat types. We measured habitat use 
by Wolves directly, and found that Wolf use of land- 
scapes was not random. We suggest that patterns of 
habitat use may be influenced by the relative avail- 
ability of different habitat types, specifically natural 
shrubs and waterways, and to a lesser degree, by re- 
cent forest cutblocks. Nevertheless, our results clearly 
show that consideration of shrub/waterway habitats is 
an important criterion for land-use decisions regard- 
ing Wolves. In our study area, Caribou prefer forests 
greater than 80 years old, especially those stands aged 
120-160 years (Szkorupa 2002), and have been found 
to avoid forest cutblocks by 1200 m (Smith et al. 
2000). As most of the winter range of these Caribou 
has been allocated for timber extraction, areas of older 
forest will become increasingly small and isolated. If 
the forest continues to be harvested at present rates, 
all Wolf packs we studied will have a substantial 
amount of logged area within their territories within 
a relatively short time. As Moose, deer and Elk are 
the primary prey of Wolves in this study area, informa- 
tion is required on how these ungulates are respond- 
ing to the changing landscape mosaic, as this may 
ultimately determine how Wolves use the landscape. 
Understanding the dynamic relationship between pred- 
ator and prey in a system undergoing rapid change 


374 


poses an enormous challenge. Future research on Wolf 
habitat use should concentrate on increasing sample 
sizes, refining habitat classifications, and linking be- 
havior with patterns of habitat use. 


Acknowledgments 

Funding for this research was provided by the West- 
Central Alberta Caribou Standing Committee, the Al- 
berta Sport, Recreation, Parks and Wildlife Foundation, 
and a University of Alberta Challenge Grant in Bio- 
diversity (supported by the Alberta Conservation Asso- 
ciation). A University of Alberta Graduate Research 
Assistantship and Margaret Brown Award in Environ- 
mental Studies and Wildlife Resources also provided 
financial assistance to G. Kuzyk. Additional financial 
support and digital forest inventory data were kindly 
provided by Weyerhaeuser Canada Ltd. and Cana- 
dian Forest Products. S. Shirkoff provided GIS sup- 
port. We are thankful for the safe and expert piloting 
conducted by C. Wilson from Bighorn Helicopters 
and D. Dennison from Coyote Air during wolf cap- 
tures and monitoring. K. Smith and R. Hayes kindly 
reviewed the manuscript and added helpful suggestions. 


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Received 4 April 2003 
Accepted 23 November 2004 


Bird Communities of the Garry Oak Habitat in Southwestern British 
Columbia 


WAYNE R. ERICKSON 


Forest Practices Branch, British Columbia Ministry of Forests, P.O. Box 9513, Stn Prov Govt, Victoria, British Columbia 
V8W 9C2 Canada 


Erickson, Wayne R. 2004. Bird communities of the Garry Oak habitat in southwestern British Columbia. Canadian Field- 
Naturalist 118(3): 376-385. 


Identifying the bird communities of a habitat could contribute to conservation efforts and provide benchmarks for ecosystem 
studies. Garry Oak (Quercus garryana) ecosystems in British Columbia are among the most endangered in Canada and 
warrant conservation. Four bird communities were determined by analyzing an extensive sample of Garry Oak habitat bird 
data. These communities were defined objectively by aggregations of the bird species themselves from across the various 
sites and areas. Characteristic species of these communities include American Goldfinch (Carduelis tristis), Spotted Towhee 
(Pipilo maculatus), Bewick’s Wren (Thryomanes bewickii), Rufous Hummingbird (Selasphorus rufus) and Chestnut-backed 
Chickadee (Poecile rufescens) in community 1; House Wren (Troglodytes aedon), Olive-sided Flycatcher (Contopus cooper), 
Purple Finch (Carpodacus purpureus), White-crowned Sparrow (Zonotrichia leucophrys), and Pine Siskin (Carduelis pinus) 
in community 2; Western Tanager (Piranga ludoviciana), Yellow-rumped Warbler (Dendroica coronata), American Goldfinch, 
Pine Siskin, Pacific-slope Flycatcher (Empidonax difficilis), Cassin’s Vireo (Vireo cassinii) and Chipping Sparrow (Spizella 
passerina) in community 3; and Northwestern Crow (Corvus caurinus) and European Starling (Sturnus vulgaris) in commu- 
nity 4. Differences between the communities are suggested from the life history traits of the species, including a community 
consisting mostly of insectivores when on breeding territory (number 1), one with species foraging primarily in shrubs and 
trees (community 3), and another with tree-nesting ground gleaners (number 4). One community (number 3) had analogues 
in two widely disparate areas: oak-associated in north-central New Mexico, and aspen (Populus tremuloides)-related in 


northcentral British Columbia; otherwise communities reported in the literature were generally not directly comparable. 


Key Words: bird communities, Garry Oak, Quercus garryana, multivariate classification, British Columbia. 


Garry Oak, Quercus garryana habitat is unique 
within Canada, and related biogeographically to Cal- 
ifornia (Erickson 1996). Garry Oak ecosystems are at 
their northern margin in British Columbia, and are 
among the most endangered ecosystems in Canada 
(Erickson 1993*, 2000a). Native stands have been 
reduced by urban development and are threatened 
with invasion by alien species. Bird communities of 
Garry Oak habitat in the Pacific Northwest have not 
yet been fully investigated. Defining these communi- 
ties is a basic step toward conservation, and a requisite 
to identifying critical habitat requirements. Predictive 
abilities and assessments for preservation and man- 
agement of the Garry Oak habitat could be strength- 
ened by further understanding these communities. 

Garry Oak occurs in an area of southwestern British 
Columbia with a distinctive modified mediterranean 
climate in the strong rain-shadow of the Olympic and 
Vancouver Island mountains. Mild, wet winters, vari- 
able precipitation and regular summer drought are typ- 
ical. The result on the landscape is a mosaic of park- 
lands with spring forb meadows and oak clumps; mossy 
bluffs often with shrub oaks; open grassy savannahs; 
and woodlands sometimes mixed with Douglas-fir 
(Pseudotsuga menziesii). As well as being diverse and 
productive for plant growth, this landscape is attractive 
for human habitation and agricultural development. 


The consequence has been habitat loss and endanger- 
ment for Garry Oak ecosystems. There is a scientific 
and conservation interest in knowing more about the 
elements of these ecosystems, including their bird 
communities. Garry Oak habitat has been selected 
as a focus of Partners in Flight for its importance to 
migrating and wintering birds in the Pacific Northwest 
(de Groot et al. 2000). Their work includes restoration 
of habitat within the British Columbia range of Garry 
Oak (Figure1). 

The purpose of this paper is to identify the bird com- 
munities of Garry Oak habitat, make comparisons of 
the life history traits of constituent species, and com- 
pare these communities with the literature on other 
habitats. 


Methods 

I used Detrended Correspondance Analysis (DCA, 
PCORD 3.0, McCune and Mefford 1997*) (Hill and 
Gauch 1980; Peet et al. 1988); interpretive graphing; 
and a quantitative similarity index (Motyka’s modifi- 
cation of the Sorenson index, Brower et al. 1990): to 
determine bird communities. 

I took records of bird species occurrence by ear and 
by sight while sampling representative ecological plots 
over 120 Garry Oak areas (Figure 2). This was part of 
an ecological reconnaissance in 1993 and 1994, cov- 


376 


2004 


in British Columbia 


| Ficure 1. Garry Oak range in British Columbia. This map is 
courtesy of British Columbia Ministry of Water, Land 
and Air Protection, December 2003. 


ering an area bounded by East Sooke, Gonzales Hill 
and Courtenay on Vancouver Island; and East Point, 
Saturna Island, and Helliwell Point, Hornby Island 
on the Gulf Islands (Erickson 1996). These locations 
are between approximately 48°N, 123°W and 49°30'N, 
125°W. The Garry Oak stands sampled were primarily 
mature, but their canopy varied from shrub-like with 
exposure, to large and open on deep soils. Elevations 
ranged from sea level to about 550 m. Plots were 
approximately 200 m?, consisted of relatively uniform 
vegetation and topography, and varied in size accord- 
ing to plant community boundaries. Species presence 
was recorded in an observation effort of approximately 
90 minutes per plot. A total of 1243 records were taken 


ERICKSON: BIRD COMMUNITIES OF GARRY OAK HABITAT 


SEH 


on 286 plots (Erickson 2000b). Sampling was primarily 
in spring, during May and June. This time interval has 
migratory bird influx, spring vegetation growth, territory 
establishment, nesting, summer plant growth cessation 
and bird dispersal. 

The data set had been previously checked with spe- 
cies accumulation curves (McCune and Mefford 1997; 
Erickson 2000b, unpublished data; Smith et al. 2002) 
to determine the adequacy of the sample. Occurrence 
by plot of all species with a frequency 5% for each 
year was entered into a data set; 23 species in 1993, and 
29 species in 1994. This qualifying criterion reduced 
the number of plots to 135 in 1993 and 127 in 1994, for 
a total of 262 in the analysis. The DCA method uses 
chi-squared distances to simultaneously ordinate, in this 
case, bird species data against plots, and vice-versa. 
Axis solutions account for, and therefore represent the 
most variation in the data set. Bird species are sepa- 
rated and referenced by their scores relative to the de- 
trended and re-scaled axes. Graphing the axis combi- 
nations provides a view of the reference coordinates 
in multivariate space. DCA is among the most widely 
used analytical methods in ecology (Peet et al. 1988). 
It has been applied in bird community work in oak 
woodland and other settings (Huff and Raley 1991; 
Pojar 1995; Garcia et al. 1998; Abernethy et al. 2001*; 
Parody et al. 2001). 

In the interpretive graphing method I framed pro- 
portional circles on Axis | vs. 2, and Axis 1 vs. 3, on 
the DCA output graphs in order to outline potential 
bird groups. The circles represent four potential bird 
groups for each year’s data, a number which was deter- 
mined from species accumulation curve results. These 


Helliwell Point 


Sea 


) Nanaimo 


Vancouver Island $ 


So 
[Sime eames | 


re; <1 UU ¢ = 
5 oe. | S 
eS oe 
Cone E 
: : Se 
Georgia Strait Ne 
Tee 
xX Sea 
2e 
NS SS x ; 
SS East Point 


se = 
Gonzales Hill 


+etoria 


Notes: 1 to 10 plots per circle. Note lateral map distortion 


\ East Sooke 


Sse 


RS ES] 


FiGure 2. Garry Oak bird community sampling on Vancouver Island, British Columbia. 


378 


TABLE |. Characteristic species classification. 


Species classification Symbol and similarity value 


d (10) 
c (7) 
n (3) 


Distinct 
Companion 
Non differential 


results indicated that groups of approximately 35 plots 
could potentially cover >60% of the total species. The 
circles were centred on the graph coordinates from 
the median of the top-ranking species. Species rank 
was judged from the scores against each axis and the 
dominant axis combination (1 and 2) in the DCA result 
(Table 1, 2). These groups quantitatively represent the 
co-occurrence of species in different plots from the 
data, and they served as the first stage in identifying 
bird communities. Species were classified according to 
their distinctness to a group from the graphs and numer- 
ic values assigned (Table 1). Quantitative similarity 
index comparisons were then completed and the groups 
aggregated into communities based on the results. 


Results 

Species scores against the DCA axes are shown in 
Tables 2, 3 and 4. Four bird communities of Garry 
Oak habitat were identified by the analysis (Figures 
3-6). Combinations of characteristic species, includ- 
ing most of the twenty top-ranking species in overall 
frequency, helped define the communities (Table 5). 
Characteristic species included American Goldfinch 
(Carduelis tristis), Spotted! Towhee (Pipilo macula- 
tus), Bewick’s Wren (Thryomanes bewickii), Rufous 
Hummingbird (Selasphorus rufus) and Chestnut-backed 
Chickadee (Poecile rufescens) (community 1); House 
Wren (Zroglodytes aedon), Olive-sided Flycatcher (Con- 
topus cooperi), Purple Finch (Carpodacus purpureus), 
White-crowned Sparrow (Zonotrichia leucophrys), and 
Pine Siskin (Carduelis pinus) (community 2); West- 
ern Tanager (Piranga ludoviciana), Yellow-rumped 
Warbler (Dendroica coronata), American Goldfinch, 
Pine Siskin, Pacific-coast Flycatcher (Empidonax dif- 
ficilis), Cassin’s' Vireo (Vireo cassinii) and Chipping 
Sparrow (Spizella passerine) (community 3- 1993 only); 
and Northwestern Crow (Corvus caurinus) and Euro- 
pean Starling (Sturnus vulgaris) (community 4- 1994 
only). 

Three groups had resulted in each of the two years, 
which were then reduced to the four communities. 
Two of these occurred across the years and two were 
unique to a single year. Communities were defined by 
the combination of their characteristic species. Simi- 
larity index values were quite low for the two across- 
year communities: 0.36 for community | and 0.43 for 
community 2. The two within-year communities had 
no similarity to each other (0 index value). 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Association with a group on both axis combinations 


unique to a group on both axis combinations 
unique to the group on one axis combination 
distinct on one axis combination 


TABLE 2. Garry Oak bird species in 1993 with the highest 
scores against DCA axes, with median species designated*. 


Species Score 
Axis |: 
Northwestern Crow 522 
Song Sparrow 499 
California Quail* 388 
Spotted Towhee* 388 
Cedar Waxwing 388 
Bewick’s Wren 378 
Axis 2: 
Cedar Waxwing 495 
Pine Siskin 446 
Bewick’s Wren 400 
Western Tanager* 388 
Cassin’s Vireo* 384 
Yellow-rumped Warbler 384 
California Quail Sl 
Song Sparrow 345 
Axis 3: 
Chestnut-backed Chickadee 428 
Rufous Hummingbird 336 
Northern Flicker* 318 
Pacific slope Flycatcher 294 
Cedar Waxwing 2S) 


TABLE 3. Garry Oak bird species in 1993 and 1994 with the 
highest scores against DCA axis combination | and 2, with 
median species designated*. 


Year and Species Score Score 
1993 axis | axis 2 
California Quail 388 371 
White-crowned Sparrow 183 250 
Olive-sided Flycatcher* 119 230 
American Goldfinch 302 342 
House Wren 69 135 
1994 

Brown Creeper 222 230 
Rufous Hummingbird 406 275 
Brown-headed Cowbird* 373 262 
Northern Flicker* 258 426 
Pacific-slope Flycatcher 216 162 
Cassin’s Vireo 254 224 


Note: The former common names (Common Flicker, Rufous- 
sided Towhee, Common Starling, and Solitary Vireo) were 
used in the data and Figures for Northern Flicker, Spotted 
Towhee, European Starling and Cassin’s Vireo. 


' The former common names, (Common Flicker, Rufous-sided Towhee, Common Starling, and Solitary Vireo) were used 
in the data and Figures 1-4 for Northern Flicker, Spotted Towhee, European Starling and Cassin’s Vireo. 


2004 


TABLE 4. Garry Oak bird species in 1994 with the highest 


scores against DCA axes, with median species designated*. 


Species Score 
Axis 1: 
Song Sparrow 583 
Northwestern Crow* 460 
European Starling* 420 
Rufous Hummingbird 406 
Axis 2: 
Common Flicker 426 
White-crowned Sparrow 40] 
Song Sparrow* 294 
Chipping Sparrow 283 
Rufous Hummingbird PAIS 
Axis 3: 
European starling 506 
Violet-green Swallow 396 
Olive-sided Flycatcher* 378 
House Wren* 358 
White-crowned Sparrow 327 
Spotted Towhee 326 


Note: The former common names (Common Flicker, Rufous- 
sided Towhee, Common Starling, and Solitary Vireo) were 
used in the data and Figures for Northern Flicker, Spotted 
Towhee, European Starling and Cassin’s Vireo. 


The overall frequency (Erickson 2000b) of these 
characteristic species gives an indication of the abun- 
dance of the communities. Community 1 was most fre- 
quent, with the #3-rank Spotted Towhee; the #4 Chest- 
nut-backed Chickadee; #6 American Goldfinch; #14 
Bewick’s Wren and #16-rank Rufous Hummingbird. 


4PiSi 


Unngmed group 


Community 3 


SomMRWa ele 


Axis | 


ERICKSON: BIRD COMMUNITIES OF GARRY OAK HABITAT 379 


Community 2 had the #5-rank White-crowned Sparrow; 
#10 House Wren; #12 Pine Siskin; #13 Olive-sided 
Flycatcher; and the #19-rank Purple Finch. Commu- 
nity 3 had the #6-rank, American Goldfinch; #7 
Pacific-coast Flycatcher; #11 Chipping Sparrow; #12 
Pine Siskin; #18 Yellow-rumped Warbler; #21 
Cassin’s Vireo; and #24-rank Western Tanager. Com- 
munity 4 had the #15 rank Northwestern Crow and the 
#26 rank European Starling. 


Discussion 
Sampling and analysis 

Communities are differentiated here by their visual 
distinctness in the multivariate space defined by bird 
frequency and composition, and by differences defined 
using thresholds in quantitative index of similarity com- 
parisons. Both measures use quantity along with com- 
position, which ensures that the communities arising 
from the data are actually different from each other. 
Although eigenvalues can measure the overall strength 
of a multivariate relationship, such as the multivariate 
coefficient of variation for each axis in an ordination, 
they are not a test statistic in Detrended Correspondance 
Analysis.This method uses detrending and rescaling to 
avoid the spurious, secondary arch effect of previous 
techniques, but this in turn prevents the use of eigen- 
values. Although the differences by year were signifi- 
cant for many species in previous t-test comparisons 
(Erickson, unpublished data), these tests are not used 
here, as they are not appropriate for analysis of non- 
experimental survey data (e.g., Hurlbert 1984). 

Sampling occurred on discrete but much smaller 
(0.02 ha) plots in this study than in many others. How- 


se ogee nl 2 
OCWa 


Community 4 
4NWCr 


FiGurE 3. Garry Oak bird communities from DCA analysis of 1993 data, Axis 1 vs. 2. Large square represents the multi- 
variate space formed from the data centred on the DCA axes. Circles are objectively defined groups or communities. 
Small square is a subjectively placed community. Bird species coordinates are marked by a cross and designated by 
the first letters of the common name (former names as noted in the text). Other data points are the plot coordinates. 


380 THE CANADIAN FIELD-NATURALIST Vol. 118 


Cc ity 4 
Unnamed group saa 


Community 3 
| ” PSEI 


aI sar 
SowRWa 


FiGure 4. Garry Oak bird communities from DCA analysis of 1993 data, Axis | vs. 3. Large square represents the multi- 
variate space formed from the data centred on the DCA axes. Circles are objectively defined groups or communities. 
Small square is a subjectively placed community. Bird species coordinates are marked by a cross and designated by 


the first letters of the common name (former names as noted in the text). Other data points are the plot coordinates. 


oF 


{ChSp 


Vie 
f 
Community 3 GS ; 
Fh OWA 
/ ie 
een ies 
| 


w 


Axis 2 


Conur a: 


4WCSp 


Community | 


94 group 4 


DSp 


Ficure 5. Garry Oak bird communities from DCA analysis of 1994 data, Axis 1 vs. 2. See explanation for Figure 4. 


ever, the number of plots was correspondingly larger 
(262 plots) and records were taken over a much longer 
observation period (90 minutes per plot) than in many 
studies. Consequently, the results do not represent ex- 
treme low values in comparison to studies using other 
methods in oak woodlands. In my species accumula- 
tion curves, about one half the species were covered 


by 15 randomly selected plots, and three-quarters of 
the species by 45 plots (Erickson 2000a). 

The average number of species detections per plot 
(3.7, 1993; 5.0, 1994: Erickson 2000a) is from the 
same order of magnitude for two count periods (3 
species, 5 species per 0.28 ha plot) in Jalisco, Mexico, 
oak woodland; but lower than two other count peri- 


2004 


Community 2 


/ 


SEI 
-HoWr | 


4GSw 


chSp  \ 


aQn \ 


4RBNu 


Axis 3 


| ; ” deel 


\ PSFI 


Axis | 


ERICKSON: BIRD COMMUNITIES OF GARRY OAK HABITAT 


\ REC'Bp 
4PuFi 


th.QCWa? 
NRG oe 
eee SvaTBCh |: 


Community3—\, Community 1 


CeWa 


381 


CoSt 


Community 4 


I 
it eet AS 


94 group 4 


BHCo \ 
4RuHu, 


FIGURE 6. Garry Oak bird communities from DCA analysis of 1994 data, Axis 1 vs. 3. See explanation for Figure 4. 


_ ods (10, 12 species: Corcuera and Butterfield 1999). 
_ My averages were much lower than those recorded on 


multiple plot, large-area studies: 21 to 25 late-spring 


_ species on five, 70 to 130 ha, stands in Oregon (Ander- 
_ son 1970), and 9 to 29 breeding species from 40 ha 


oak census plots in Pennsylvania (Probst 1980). My 
total number of species detections (66) is higher than 
in a number of studies in oak woodland (e.g., 38 spe- 
cies in Dedon et al. 1984; 20 to 62 species in Leidolf 
et al. 2000; 43 to 51 species in Stone no date*; 50 and 
58 species in each of two seasons, five forest types, 
Corcuera and Butterfield 1999) but is lower than in 
some other comparable results (e.g., 77 species in Gar- 
cia et al. 1998). My total would be increased by an 
additional 12 species which I recorded as “out of plot’, 
“out of habitat’, or “overhead”. However, much of this 
difference is irrelevant, in that my interest in defining 
bird communities was at the plot (not area) level, and 
was focused on frequently occurring species which are 
easily detected within my plot parameters, not on less 
common species used to round out a full species list. 
The graphical location method is an objective one, 
in its centering technique and the proportional alloca- 
tion of multivariate space to each community. Unbi- 
ased shapes, such as squares or circles, were used to 
define multivariate species aggregations and overlap 
in composition was accepted. Other methods include 
subjective ones (e.g., Pojar 1995), and other objective 
approaches, such as the use of a “fuzzy-clustering 
partition coefficient” (Abernethy et al. 2001*). These 
methods tend to result in elliptic forms, irregular bound- 
aries and volumes of multivariate space, and no overlap 
in composition. The elliptical forms assume a trend in 
dimensionality that does not actually occur with the 


DCA methodology, as it has demonstrated potential 
to produce evenly distibuted sample points in multi- 
variate space (Peet et al. 1988). 


Communities ; 

Of the six groups first resulting from the DCA analy- 
sis, two from each year were similar enough (similarity 
index > 0.33) to combine. One group from 1994 was 
dropped because it had less than my conceptual thresh- 
old of three species. Two within-year communities were 
not differentiated by the results of the other year, but 
it was possible to represent them subjectively. To do so, 
I placed the square frame (Figures 3-6) on the graphs of 
the other year, and in doing so, included species and 
multivariate space not already covered. Through this 
process, Dark-eyed Junco (Junco hyemalis) was added 
to community 3 as a distinct species in 1994, although 
in 1993 it was limited to one of the two one-axis com- 
binations. The two top-ranking species in frequency, 
Orange-crowned Warbler (Vermivora celata) and Amer- 
ican Robin (Turdus migratorius) did not contribute to 
the characteristic combination of species for the com- 
munities. American Robin did qualify as a character- 
istic species, but was not used because it was in three 
out of four communities. 

Consideration was given to dropping another group 
which occurred only in 1994, as it had only two dis- 
tinct species, Northwestern Crow and European Star- 
ling. However, it was kept (as Community 4) because 
these two species are potentially important as ecolog- 
ical indicators. They may both signify a disturbance 
zone and represent a source of disturbance themselves, 
via nest predation and nest cavity competition. In 
addition, this community can be characterized by the 
absence of other species. There may be an inversely 


382 


THE CANADIAN FIELD-NATURALIST 


TABLE 5. Bird communities and characteristic species of Garry Oak habitat. 


Species Group 


94-3 


Community | 
American Goldfinch 
Spotted Towhee 
Bewick’s Wren 
Rufous Hummingbird 
Chestnut-backed Chickadee 


Community 2 
House Wren 
Olive-sided Flycatcher 
Purple Finch 
White-crowned Sparrow 
Pine Siskin 


Community 3 
Western Tanager 
Yellow-rumped Warbler 
Pacific-slope Flycatcher 
Cassin’s Vireo 
Chipping Sparrow 
Dark-eyed Junco* 


Community 4 
European Starling 
Northwestern Crow 


Other species: 


Song Sparrow n 

California Quail d 

Violet-green Swallow 

Orange-crowned Warbler n 


Townsend’s Warbler 
Brown Creeper 


Cedar Waxwing n 
Swainson’s Thrush n 
Common Flicker 

American Robin n n 


Vol. 118 
93-3 94-2 93-2 94-1 
Cc (¢ 
(e n 
n 
n nN 
d d 
Cc d 
(e Cc n 
Cc n n 
n n (© 
d d 
d 
n Cc 
(Gj 
c c 
n 
d 
d 
(& 
n 
n 
n n 
n n 
n 
Nn (© Cc 


Notes: Communities are shown in the boxes. See Table | for species classification codes. The addition of Dark-eyed Junco 


to community 3 is partly subjectively derived. 


proportional relationship to the presence of North- 
western Crow and European Starling across various 
sites. In this study, the lack of importance of another 
disturbance indicator, Brown-headed Cowbird (Molo- 
thrus ater), contrasts with the high numbers of cowbird 
nestlings on two Garry Oak sites west of Victoria, 
British Columbia (Shepard 2000). However, the present 
results reflect only detections of adults and fledged 
birds in a reconnaissance investigation. 

Cross-year similarity index values were relatively 
low, approximately one-half of my expected threshold 
value of 0.66. Therefore I considered using the concept 
of assemblages to denote a weaker level of association. 
However, both the terms community (e.g., Huff and 
Raley 1991; Pojar 1995; Garcia et al. 1998; Abernethy 
et al. 2001*; Parody et al. 2001) and assemblage (e.g., 


Manuwal 1986; Corcuera and Butterfield 1999; Hagar 
and Stern 2001) receive only general use in the litera- 
ture, and the term community is referred to more wide- 
ly. The year differences I encountered could relate to 
increases in bird frequency and dominance shifts in 
1994, possibly linked to yearly weather (Haila et al. 
1993; Erickson 2000b). 

The communities may differ in the life history traits 
of their characteristic species, as shown in Table 6 
(Leidolf et al. 2000; Ehrlich et al. 1988), and there- 
fore in their general habitat and guild use. Species of 
community | have mixed nest locations and most are 
insectivorous when on breedng territory. Those of com- 
munity 2 have an equal dominance of granivores with 
insectivores. Most species of community 3 forage in 
shrubs and trees. Community 4 species are tree-nest- 


2004 


Community and species 


Community 1: 


Nest Location 


TABLE 6. Life history traits for characteristic species of Garry Oak bird communities. 


Foraging Layer 


ERICKSON: BIRD COMMUNITIES OF GARRY OAK HABITAT 


American Goldfinch SH 
Spotted Towhee GR/ SH 
Bewick’s Wren TR 
Rufous Hummingbird VN/TR 
Chestnut-backed Chickadee SN 
American Robin SH/ TR 
Community 2: 
House Wren TR 
Olive-sided Flycatcher TR 
Purple Finch TR 
American Robin SH/ TR 
White-crowned Sparrow SH 
Pine Siskin TR 
Community 3: 
Western Tanager TR 
Yellow-rumped Warbler TR 
American Goldfinch SH 
American Robin SH/ TR 
Pine Siskin TR 
Pacific-slope Flycatcher TR 
Cassin’s Vireo TR 
Chipping Sparrow SH/ TR 
Dark-eyed Junco GR 
Community 4: 
Northwestern Crow TR 
European Starling TR 


Foraging Method Food Type 
HB/ SH/ TR FG GV 
GR GG IN/ OM 
GR GG IN 
HB HG NE 
SH/ TR BG IN 
GR/ SH/ TR GG/FG IN/ FR 
GR GG IN 
AIR HA IN 
GR GG GV 
GR/ SH/ TR GG/ FG IN/ FR 
GR GG GV 
HB/ SH/ TR FG GV 
SH/ TR FG IN 
SH/ TR FG IN 
HB/ SH/ TR FG GV 
GR/ SH/ TR GG/FG IN/ FR 
HB/ SH/ TR FG GV 
AIR HA IN 
SH/ TR FG IN 
GR GG IN/ GV 
GR GG GV 
GR GG OM 
GR GG IN 


Note: Nest location: TR-tree, VN-vine; GR-ground, SH-shrub, SN-snag. Foraging Layer: AIR-air, GR-ground, HB-herb, SH- 
shrub, TR-tree. Foraging Method: BG-bark glean, FG-foliage glean, GG-ground glean, HA-hawk, HG-hover and glean. Food 
Type: FR-frugivore; GV-granivore; IN-insectivore; NE-nectarivore, OM-omnivore. 


ing ground gleaners. Shared traits are as follows: tree 
nesting insectivores (community 2,3) and ground glean- 
ing (1,2,4). The comparison may suggest general habi- 
tat and guild relationships for the communities. This 
could include open savanna settings (community 2,4) 
and shrubby woodlands or shrub oak (community 
1,3). More work is needed on habitat relationships 
though. 


Comparison with other studies 

In this study, bird communities have been delineat- 
ed within the Garry Oak habitat at a detailed level by 
aggregations of the bird species themselves. This ap- 
proach is not typical in the literature. There are indica- 
tions of different bird communities in the discussion 
of Hagar and Stern (2001) for Garry Oak woodlands, 
including Chipping Sparrow as a species of semi-open 
woodland and Orange-crowned Warbler as shrub- 
associated. Dedon et al. (1984) employed habitat suit- 
ability class lists, possibly comparable to more detailed 
communities, in California Black Oak (Quercus kel- 
loggii) habitat. One shrub community was identified 
to subdivide Gambel Oak (Quercus gambellii) wood- 
lands (Stone no date*) in Colorado. Artman et al. 
(2001) found in Ohio that differences in bird commu- 
nities within mixed oak forests could be related to 


three moisture index classes. Pojar (1995) described 
separate bird communities within Trembling Aspen 
forests of north-central British Columbia. 

In most oak woodland bird studies, bird communities 
have been identified at a more general level. Broad 
habitat groupings have formed bird communities, 
guilds have been separated by shared traits, or com- 
munities have been grouped by successional stages 
(e.g., Anderson 1970; Smith 1977; Probst 1980; Davis 
et al. 2000; Hagar and Stern 2001). Both Leidolf et al 
(2000) and Stone (no date*) described bird commu- 
nities associated with four different Gambel Oak habi- 
tats (i.e., oak woodland, submontane shrub, Ponderosa 
Pine (Pinus ponderosa)/oak woodland, and mixed 
conifer/oak woodland), and locations. Garcia et al. 
(1998) and Corcuera and Butterfield (1999) referred 
to separate bird communities in oak woodlands, mixed 
oaks (Quercus rugosa, Q. candicans, Q. obtusata, Q. 
laurina) and other groupings for north-central Michoa- 
can, Mexico. 

Some comparisons and comments can be made 
about the composition of communities from the present 
study in relation to these others. Shepard (2000) out- 
lined the importance of House Wren (found in com- 
munity 2) for two Garry Oak sites west of Victoria, 


384 


British Columbia. However, those sites had steady num- 
bers of Red-breasted Nuthatch (Sitta canadensis) and 
Brown Creeper (Certhia americana), species which 
were unimportant in the present study. Most of the 
frequent birds from my study were present in Black 
Oak habitat in northern California, but no more than 
one or two species were listed by habitat suitability 
class (Dedon et al. 1984). Five of the seven species 
of community 3 were resident or breeding species in 
the ponderosa pine/Gambel Oak forest in north-central 
New Mexico (Leidolf et al 2000). The two remaining 
species are not found in the range of Gambel Oak. 
However, the shrub subdivision in Gambel Oak bird 
communities (Stone no date*) is not similar to the 
four communities except for the presence of Spotted 
Towhee. A community from Trembling Aspen forests 
in north-central British Columbia had similarities 
(4/7 species) to community 3 (Pojar 1995). Although 
this was labelled a group of conifer-related species, 
three of four species were associated with Gambel Oak 
and Trembling Aspen in southwestern Colorado (Stone 
no date*). In addition, Shepard (2000) mentioned high 
densities of Pacific-slope Flycatcher, also a species of 
community 3; and the importance of Chesnut-backed 
Chickadee (found in community 1), for two Garry Oak 
sites in British Columbia. These species have similarly 
been thought of as conifer (Douglas-fir)-related (Huff 
and Raley 1991). Neither the other two Trembling 
Aspen bird communities (Pojar 1995), nor the ones 
described for Douglas-fir habitat (Shepard 2000; Huff 
and Raley 1991) had any substantial similarity to the 
four communities in the present study. 


Summary 

Testing of wildlife habitat relationship models based 
on broad habitats (e.g., oak woodland, old-growth 
Douglas-fir forest) has given results which questioned 
their basic utility (e.g., Dedon et al. 1984; Laymon 
1989). This suggests that other methods could be 
examined. An alternative approach, which empirical- 
ly defines spatial and temporal bird aggregations as 
bird communities, is presented here, and may war- 
rant further investigation. 

The four communities identified in this work could 
serve as benchmarks in Garry Oak ecosystem studies. 
Like any classification, they should be tested with fur- 
ther field work and analysis, particularly for differen- 
tiation by habitat features. For example, Erickson and 
Campbell (2001) related correlations involving mois- 
ture regime, oak diameters, tree form complexity, total 
wildlife habitat features, and, for one year, average 
number of bird species per plot. Monitoring is recom- 
mended as a first step in maintaining characteristic 
Garry Oak bird communities, a focus which should 
parallel growing concern for this endangered habitat 
and contribute to increasing conservation efforts. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Acknowledgments 

The field work for this study was completed in an 
M.Sc. program at the University of Victoria, toward 
which British Columbia Ministry of Forests approved 
a part-time education leave. Both Canadian Wildlife 
Service and a Canadian Wildlife Service employee’s 
fund provided grants which assisted in the first year 
of the study. I thank A. J. Erskine for a helpful 
review. 


Documents Cited (marked * in text) 

Abernethy, V. J., D. I. McCracken, A. Adam, I. Downie, 
G. N. Foster, R. W. Furness, K. J. Murphy, I. Ribera, 
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McCune, B., and M. J. Mefford. 1997. PCORD: Multivariate 
analysis of ecological data, version 3.0. MjM Software 
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Stone, E. no date. Avian communities of Gambel oak habitats 
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Februrary 2003): www.colorado.edu/epob/epob4630estone/ 
ES/ResearchStuff/Oak/Oak. html 


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Received 17 April 2003 
Accepted 24 January 2005 


Invertebrate Diversity under Artificial Cover in Relation to Boreal 
Forest Habitat Characteristics 


STEVEN H. FERGUSON! and DANIELLE K. A. BERUBE? 


'Fisheries and Oceans Canada, Freshwater Institute, 501 University Crescent, Winnipeg, Manitoba R3T 2N6 Canada 
> Department of Biology, Lakehead University, 955 Oliver Road, Thunder Bay, Ontario P7B 5E1 Canada 


Ferguson, Steven H., and Danielle K. A. Berube. 2004. Invertebrate diversity under artificial cover in relation to boreal forest 
habitat characteristics. Canadian Field-Naturalist 118(3): 386-392 


We investigated invertebrate diversity in boreal forests using an experimental design that consisted of counting soil inverte- 
brates under artificial cover. The aim was to assess the utility of using soil invertebrate diversity as a measure of ecosystem 
health. The study area was grouped into five habitats: upland hardwood, lowland hardwood, conifer, shrub, and conifer- 
grass. Simpson’s and Shannon’s indices of invertebrate diversity were negatively correlated with percent herbaceous cover. 
Number of recognizable taxonomic units (RTU richness) was negatively correlated with percent litter cover. The number of 
individual invertebrates was positively correlated with soil moisture and negatively correlated with percent conifer cover. 
Invertebrate diversity varied among habitat types, with conifer forests (spruce, fir, pine) having the highest diversity and 
regenerating conifer-grass forests having the lowest diversity, suggesting that successional stages affect diversity. The most 
productive sites, upland and lowland hardwood habitats, had the highest abundance of soil invertebrates, although interme- 
diate diversity compared to the other five habitats. The results are consistent with the view that diversity increases and then 
decreases with productivity and disturbance over succession (ca. 50-100 yr). Hence, maintenance of soil invertebrate diversity 


in managed boreal forests requires the provision of a varied landscape with a mosaic of disturbance regimes. 


Key Words: arthropods, biodiversity, conifer, earthworms, indices of diversity, moisture, Ontario. 


Concerns about the effects of the widespread loss 
of biodiversity have prompted many recent studies 
investigating the relationship between biodiversity 
and ecosystem function (Symstad et al. 2000). Inver- 
tebrates are important in the functioning of nearly all 
environments, with changes in species composition 
potentially reflecting changes in the ecosystem (Majer 
1990; Madden and Fox 1997). Hence invertebrates 
are increasingly being viewed as reliable indicators to 
assess human impacts on the general level of distur- 
bance of an ecosystem (Majer 1983; Greenslade 1984; 
Andersen 1990). Biodiversity includes all levels of 
natural variation and thus diversity indices provide a 
relative measure of variation within a community (Til- 
man and Pacala 1993). Monitoring diversity across 
spatial and temporal scales allows for measurement of 
system complexity, functionality, and stability. Knowl- 
edge of diversity helps in understanding changes in 
ecosystem complexity before and after disturbance. 
Information on the habitat characteristics that influence 
diversity at various levels and knowledge of habitat 
changes resulting from human disturbance are required 
for management and conservation (Madden and Fox 
19977): 

Forest managers can assess diversity changes asso- 
ciated with human disturbances that include various 
forestry practices through an understanding of the 
relationship between animal diversity and forest struc- 
ture to determine ecosystem changes (Noss 2000). 
Animal diversity includes soil invertebrates common- 


ly found under logs and rocks in managed forests 
(Kolstrom and Lumatjarvi 1999). Downed wood is im- 
portant for organisms in providing shelter and moisture, 
and in preventing light penetration. The use of soil 
fauna diversity has the potential to act as a surrogate of 
forest biodiversity. A number of forest characteristics 
have been shown to relate to soil invertebrate diversity, 
including understorey vegetation and litter (Bird et al. 
2000), plant functional diversity (Siemann et al. 1998), 
coarse woody debris (Marra and Edmonds 1998), 
conifer species composition (Lattin 1993), forest suc- 
cession (Paquin and Corderre 1997), soil moisture 
(MacKay et al. 1986) and structural complexity (Fer- 
guson 2001). 

To further this research, we used a method of sur- 
veying invertebrate diversity under artificial cover and 
relate indices of invertebrate diversity to measured for- 
est characteristics. The study design consisted of (1) 
surveys of soil invertebrates (springtails, beetles, cen- 
tipedes, slugs, earthworms and isopods) found under 
sand-filled cardboard boxes placed on the forest floor; 
and (2) surveys of the sampled forest characteristics 
(snags, logs, soil moisture, overstorey, understorey, 
and ground cover). Forest characteristics were used 
to identify habitats within a boreal forest landscape, 
and diversity of soil invertebrates was measured. The 
goal was to relate forest habitat characteristics to soil 
invertebrate diversity to assess their utility as surro- 
gate measures of ecosystem changes associated with 
forest management practices. 


386 


2004. FERGUSON and BERUBE: INVERTEBRATE DIVERSITY 327 


RS 
1.0 

cb) 

O 

S 

is) 

—~ 

jp 

<2) 

© 

0) 0.5 

iS 

rs) 

=) 

LL 


Forest types 


Ficure 1. Location of study area in northwestern Ontario and dendrogram of 20 plots (0) located within a 1 km? stand of 
boreal forest grouped using cluster analysis based on forest characteristics (e.g., Overstorey. understorey, ground 
cover, moisture, coarse-woody debris). 


388 


Study Area 

The study area (Figure 1) was located in northwest- 
ern Ontario, Canada, and consisted of a 1 km? area of 
boreal forest located along the McIntyre River within 
Lakehead University’s natural forest (48°22'N, 
89°19'W). The mixed boreal forest consisted of Jack 
Pine (Pinus banksiana Lamb.), Black Spruce (Picea 
mariana (Mill.) B.S.P.), Balsam Fir (Abies balsamea 
(L.) Mill.), White Spruce (P. glauca (Moench) Voss), 
White Birch (Betula papyrifera Marsh.), and Trem- 
bling Aspen (Populus tremuloides Michx.). The study 
site lies within the Boreal Ecosystem that consists of 
rolling rocky uplands with coarse well-drained soils 
(Rowe 1972). The climate is humid continental with 
a mean minimum January air temperature of -15°C 
and a mean maximum daily air temperature for July 
of 18°C (Environment Canada 2001"). Mean annual 
precipitation is approximately 700 mm, including a 
mean winter snowfall of 196 cm (Environment Cana- 
da 2001"). For the study period (May to September), 
mean monthly daily air temperature (1961-1990 nor- 
mals) varied from 9.0°C in May to 17.7°C in July 
(annual mean = 2.4°C) and precipitation varied from 
69.3 mm in May to 88.5 mm in August (annual mean 
= 58.6 mm) (Environment Canada 2001"). 


Methods and Materials 
Study design and sample plots 

The sampling method was designed to assess soil 
invertebrate diversity under a standardized collection 
technique that optimized sampling replication, effort, 
and coarse taxonomic resolution. Coarse woody debris 
occurred in various shapes, sizes, and material. The 
use of cardboard boxes standardized the collection and 
reduced this variability while sampling a more diverse 
fauna than other methods such as pitfall traps. We 
decided to use a coarse taxonomic resolution (Bolger 
et al. 2000), thereby allowing inexperienced observers 
with minimal training to obtain reasonable survey 
counts efficiently. A trade-off associated with group- 
ing invertebrates occurs between ease of surveys by 
observers with minimal taxonomic experience and 
the loss of more detailed guild and life history infor- 
mation related to individual species. 

Twenty plots, each consisting of three adjacent boxes, 
were randomly distributed within the forest (minimum 
distance between plots was 5 m; Figure 1). Each box 
consisted of two 2-liter milk cartons with a plastic 
coating of red and white color. These were fitted one 
inside the other to create a solid box and filled with 
sand (approximately 2 kg) to create a footprint-sized 
depression 21 by 9.5 cm. Boxes depressed the leaf 
litter an average of 1.3 cm (Ferguson 2000), creating 
a microhabitat of increased humidity and decreased 
temperature similar to that beneath a log or rock rest- 
ing on the forest floor. 

Boxes were overturned and the numbers of all soil 
invertebrates (>1 mm) were visually counted. Boxes 
were lifted individually without disturbing adjacent 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


boxes or the underlying litter. Twenty weekly surveys 
were conducted from 9 May to 24 September 2000 
between the hours of 1100 to 1700. The survey num- 
bers reflect a relative abundance of soil invertebrates. 
Observer bias was consistent across the landscape. 
Recognizable Taxonomic Units (RTU; Bolger et al. 
2000; Ferguson 2004) were used to group all macroin- 
vertebrates (>3 mm) observed under boxes based on 
differences in size and feeding habits (Eisenbeis and 
Wichard 1987; Brock et al., 1994) and included spring- 
tails (Collembola), spiders (Araneae), ants (Formicidae), 
ant larvae, centipedes (Chilopoda), Diplura, adult flies 
(Diptera), phytophagous mites (Acarina: Oribatida), 
bugs (Hemiptera and Homoptera), Pseudoscorpionida, 
wasps (non-ant Hymenoptera), moth and butterfly larvae 
(Lepidoptera), Gastropoda (snails and slugs), Isopoda 
(woodlouse; Tracheoniscus rathkei), beetles (Coleoptera 
and Staphylinidae species — adults and larvae), milli- 
pedes (Diplopoda), and earthworms (Oligochaeta). 


Forest characteristics measured 

Site-specific habitat variables were measured 21- 
23 August 2000 using 5 x 5 m quadrats centered at 
three box plots. One quadrat was located at the centre 
of each plot. Four other quadrats were located 10 m 
from the centre in cardinal compass directions. A total 
of 100 quadrats were sampled for the following habitat 
characteristics: percent overstorey cover (>5 m), per- 
cent understorey cover (saplings and shrubs 0.5-5 m), 
percent herbaceous cover, percent litter cover, percent 
grass cover, percent moss cover, percent fern cover, 
number of snags (dead standing trees with dbh > 5 cm), 
number of decaying logs (> 5 cm diameter), percent 
cover by conifers, and a relative measure of soil mois- 
ture (1.e., xeric=1, mesic=2, hydric=3). Values for the 
five quadrats were averaged for each plot. 


Measures of invertebrate diversity 

Diversity of soil invertebrates was calculated for 
each plot across surveys using RTU richness (num- 
ber of RTU at each site), Shannon-Wiener, and Simp- 
son’s Indices of Diversity (Ludwig and Reynolds 
1988). Shannon’s entropy (H) is a measure of species 
diversity used in relation to relative frequencies 
(probabilities) of the different species i of the sample 
and was calculated as: 

H=-% [(n/n) In(n/n)] , from i=1 ton (1) 
where n, is the number of individuals belonging to 
the ith RTU in the sample and 7 is the total number 
of individuals in the sample. H = 0 (minimum value) 
when the sample contains only a single RTU, where- 
as diversity H increases with the number of RTUs. H 
is maximum when all RTUs are equally distributed in 
the sample. The Shannon-Wiener index of diversity 
is sensitive to changes in the rare species in a com- 
munity sample (Pielou 1966). Invertebrate diversity 
was also measured by Simpson’s index of diversity 
(Simpson 1949), which is sensitive to changes in the 
more abundant species and was measured as: 


A= 1- Yn (n, - 1)/n(n-1) (2) 


2004 


Simpson’s index varies from 0 to 1, and gives the 
probability that two individuals drawn at random from 
a population belong to the same RTU. If the number 
approaches () then individuals belong to the same RTU 
and the diversity of the community sample is low. 
These diversity measures were calculated using the 
minimum number of invertebrates observed for the 
RTU in a given sampling period and were represent- 
ed by an average value for each survey. 


Statistical analysis 

We performed a cluster analysis of the 11 forest 
measures to group the 20 sampling sites into forest 
habitats. Classification of habitats types was conducted 
by the average-linkage clustering method using a Euclid- 
ean distance similarity index (Romesburg 1984). All 
variables were standardized between 0 and 1. 

Many (8 of 17) forest and soil invertebrate variables 
were not normally distributed (Wilk’s statistic) and 
transformations (e.g., log, arc-sine) failed to normalize 
all variables. Therefore, we used nonparametric analy- 
ses by ranking nonparametric data before correlation 
analyses (Conover and Iman 1981). Measures of diver- 
sity (Simpson’s and Shannon’s), RTU richness, and 
number of individuals were normally distributed and 
did not require transformations. We report untrans- 
formed means in the Figures and Tables in the Results 
section to simplify presentations. 

Analyses were performed to determine the relation- 
ship between (1) soil invertebrate diversity indices and 
forest characteristics, and (2) soil invertebrate numbers 
for each of the abundant RTU (springtails, beetles, 
centipedes, slugs, earthworms, and isopods) and for- 
est characteristics. We tested for significant effects of 
forest characteristics on dependent measures (e.g., 
diversity) with Spearman’s correlations and partial 
correlation analyses (i.e., multiple regression of ranked 
data without replacement). ANOVA was used to com- 
pare forest characteristics, indices of diversity, and soil 
invertebrate abundance relative to forest types, fol- 


FERGUSON and BERUBE: INVERTEBRATE DIVERSITY 389 


lowed by a Tukey multiple range test if the ANOVA 
was significant. Spearman correlations for nonpara 
metric data and Pearson’s correlations for parametric 
data compared RTU abundance with forest character- 
istics. Sample units were the 3-box groups (n = 20) 
sampled every week (n = 20). All statistical analyses 
were done using SAS (SAS Institute Inc., Cary, North 
Carolina 1987) statistical software for microcomput- 
ers: 


Results 

Both indices of soil invertebrate diversity (Shan- 
non’s and Simpson’s) were negatively related to per- 
cent herbaceous cover (R? = 0.32, F = 5.89, P = 0.01, 
n= 20 and R? = 0.35, F = 6.64, P = 0.01 respectively; 
Table 1, Figures 2A, and 2B). RTU richness (number 
of RTU) was negatively correlated with percent litter 
cover, which explained 48.6% of the variance in the 
model (F' = 17.0, P= 0.001, n = 20; Table 1, Figure 2C). 
Diversity indices were also related to litter cover as 
herbaceous cover and litter cover covaried (r = -0.94). 
Soil moisture (69.5% of variation explained) and per- 
cent conifer cover (9%; Figure 2D) best explained the 
number of individual invertebrates. Greater numbers 
of soil invertebrates occurred in wet habitats and with 
less conifer cover (Table 1). 

The forest measures were grouped into five forest 
types defined as upland hardwood, lowland hard- 
wood (mesic), shrub, conifer, and conifer-grass (Fig- 
ure 1). The upland hardwood forest consisted prima- 
rily of Trembling Aspen, with little understorey (47% 
cover) and the most abundant herbaceous cover 
(90%; Table 2). The soil invertebrate community in 
the upland hardwood forest consisted of numerous 
individuals (mean = 24/plot), moderate RTU diversity, 
and large numbers of isopods (Table 2). The lowland 
hardwood habitat consisted of mixed hardwoods in a 
mesic site that included an intermittent stream with a 
deep humus layer. The soil invertebrate community 


TABLE 1. Seven multiple regression results used to determine the significant effects of 11 forest characteristics on (1) Shannon’s 
index of diversity; (2) Simpson’s index of diversity; (3) RTU richness (where RTU = recognizable taxonomic units); (4) 
Number of individual invertebrates, (5) springtail abundance; (6) earthworm abundance; (7) isopod abundance. Of the 11 
explanatory variables included (soil moisture, % overstorey, % understorey; % herb cover; % litter; % grass: % moss; % ferns: 
number of snags; number of logs; and % conifer cover), only those with significant (P < 0.05) relationships are shown. 


Dependent Independent Coefficient Partials Model 
variable variable direction R? R? P 
(1) Shannon’s diversity index % herb cover negative 0.318 0.318 0.008 
(2) Simpson’s diversity index % herb cover negative 0.349 0.349 0.006 
(3) RTU richness % litter cover negative 0.486 0.486 0.001 
(4) Number of RTU Soil moisture positive 0.695 0.695 0.0001 
% conifer cover negative 0.087 0.782 0.02 
(5) Springtail abundance % conifer cover negative 0.224 0.224 0.04 
(6) Earthworm abundance Conifer cover negative 0.657 0.657 0.0001 
Number of logs positive 0.100 0.757 0.02 
(7) Isopod abundance % conifer cover negative 0.624 0.624 0.0001 


390 


Shannon's index 


0 25 50 75 
Percent herb cover 


Species richness 


100 


0 25 50 75 
Percent litter cover 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


© 
Ce) 


Simpson's index 


Percent herb cover 


800 


600 


400 


Number of individuals 


0 25 50 75 
Percent conifer cover 


FIGURE 2. Bivariate relationships between two indices of diversity (Simpson’s and Shannon’s) of soil invertebrates and major 
explanatory habitat variables: (A) Shannon’s index of diversity, and percent herbaceous cover (y = -0.307x + 2.29, 
7 = 0.318); (B) Simpson’s index of diversity and percent herbaceous cover (y = -0.0810x + 0.883, 7 = 0.349); 
(C) RTU richness (number of Recognizable Taxonomic Units) and percent litter cover (y = -0.0692x + 18.6, 7° = 0.486): 
and (D) number of individual invertebrates and percent conifer cover (y = -9.38x + 712, r? = 0.695). 


living in this forest consisted of moderate diversity, 
the greatest number of individuals (mean = 37/plot), 
and the most springtails and earthworms. The shrub 
habitat had the highest understorey cover (78%), low- 
est overstorey (34%), and moderate conifer and litter 
cover. The soil community in the shrub habitat was 
moderately diverse and consisted of a moderate num- 
ber of individuals (mean = 16/plot). The conifer for- 
est was the most extensive habitat and was composed 
of a mixed softwood/ hardwood (56/44%) forest with 
high overstorey cover (66%), high understorey cover 
(66%), low herb cover (33%), and high litter cover 


(64%). The conifer forest supported the highest soil 
invertebrate diversity although few RTU (low RTU 
richness) and few individuals (mean = 10/plot; Table 
2). The conifer-grass forest type was represented by a 
relatively low overstorey (55%), intermediate under- 
storey (56%), high conifer cover (53%) with the most 
grass (13%) and more herb than litter cover (66/22%). 
The soil invertebrate community in the conifer-grass 
habitat was the least diverse with the lowest RTU 
richness and fewest individuals per plot (n = 9). 

The abundance of three out of the six most numerous 
RTU were significantly related to forest characteristics 


2004 FERGUSON and BERUBE: INVERTEBRATE DIVERSITY 39 | 
TABLE 2. Comparison of forest and soil invertebrate characteristics for five forest types. Mean invertebrate abundances 
(numbers per m7) are given for six of the more common invertebrates. Means with the same letters do not differ significantly 


according to Tukey’s multiple range test. 


Upland 
hardwood 

Variable F IP (n= 6) 
Forest characteristics 
Snags 6.0 0.005 ab 13.3 
Logs 10.5 0.0003 ab 7.0 
Soil moisture 26.7 0.0001 b 0.6 
Overstorey Sel 0.008 ab 52.8 
Understorey 4.4 0.01 b 46.5 
Herb 28.3 0.0001 a 89.5 
Litter 26.0 0.0001 c 2.8 
Grass 2967]: 0.0001 10), 117/ 
Conifer 19.0 0.0001 b 21.8 
Indices of diversity 
Shannon’s 5.64 0.006 a2.15 
Simpson’s 37 0.002 ab 0.191 
RTU richness S/3 0.005 a 19.3 
Number of individuals 9.11 0.0006 ab 475 
Soil invertebrates 
Springtails 3.64 0.03 ab 67.8 
Beetles 0.6 0.70 16.5 
Centipedes 2 0.34 US) 
Slugs 0.8 0.52 4.0 
Earthworms 8.8 0.002 a 54.5 
Isopods ae) 0.005 a 127.3 


(Table 1). Springtail abundance increased with greater 
litter cover (22.4% of variation explained; F = 5.2, 
P =0.04). Earthworm abundance was negatively cor- 
related with percent conifer cover (65.7% explained 
variance) and positively correlated with number of logs 
(10%; F = 34.4, P < 0.001). Isopod abundance was 
negatively related to percent conifer cover (62.4% of 
variation explained; F = 29.9, P < 0.001; Table 1). In 
contrast, beetles, centipedes, and slugs were relatively 
evenly distributed across forest types and their abun- 
dance was not significantly related to forest charac- 
teristics (not shown in Table 1). 


Discussion 

Our findings are consistent with the view that diver- 
sity is greater in the most common habitat within a 
varied landscape, such as a mixed-wood boreal forest 
(<80 y) and is often characterised by intermediate 
productivity and disturbance. Simpson’s and Shannon’s 
Indices of Diversity explained similar amounts of vari- 
ation in the pattern of diversity of soil invertebrates 
with forest characteristics. Both found percent herba- 
ceous cover the most important environmental factor 
explaining the pattern of soil invertebrate diversity 
among forest habitats. Also, both indices showed simi- 
lar correlations, in magnitude and direction, such that 


Forest types 
Lowland Shrub Conifer Conifer- 
hardwood grass 
(n = 2) (n=4) (n=6) (n= 2) 
a 16.0 b 4.3 ab 7.5 b 5.5 
a 15.0 o23 be 2.7 c 2.0 
a 1.4 c 0.0 c 0.1 c 0.0 
a 69.5 b 34.3 a 65.7 ab 54.5 
ab 54.5 a 77.8 ab 66.3 ab 56.0 
ab 68.5 be 56.8 c 32.8 b 65.5 
cb 14.0 b 31.8 a 64.2 cb 21.5 
b 0.5 b 3.5 bee, a 13.0 
b 26.0 ab 36.0 a 55.8 a 53.0 
ab 1.99 ads Lif; a 2.19 b 1.90 
b 0.155 b 0.147 b 0.142 a 0.228 
ab 17.0 ab 16.3 ab 14.5 b 14.0 
a 723 be 313 c 190 c 179 
a 120.5 ab 65.0 b 46.2 ab 76.5 
10.0 eS, 16.5 4.5 
10.5 12.0 6.5 20.0 
6.0 3:3 6.3 6.0 
a 116.5 ab 32.3 b 21.0 b 14.0 
ab 96.5 ab 64.5 b 25.3 b 14.5 


as diversity increased the percentage of herbaceous 
cover decreased. In contrast to our findings, soil inver- 
tebrates from 27 orders did not differ significantly 
among five different landscapes (with the exception 
of earthworms) (Kalisz and Powell 2000). 

We found a negative relation of soil invertebrate 
diversity with increasing composition of conifer veg- 
etation. Few herbs and more litter are present under 
the conifer cover due to greater soil acidity (Kimmins 
1997). Numbers of invertebrates were greater under 
herbaceous cover but diversity was reduced. Other 
studies have found invertebrates occurring in greater 
numbers under deciduous and herbaceous cover rela- 
tive to conifer litter (Wallwork 1983; Paquin and 
Coderre 1997; Hammond 1997; Marra and Edmonds 
1998). Within varied forest landscapes, the greatest 
diversity of soil invertebrates likely occurs for the most 
common (in time and space) successional community. 
In northern boreal forests this is likely for older conifer 
stands with greater overstorey and reduced ground veg- 
etation understorey. Removal of conifers by natural 
disturbance or forest harvesting likely results in lower 
initial diversity with increasing diversity over time as 
other plants colonize the habitat (i.e., succession). We 
found greater invertebrate density under herbaceous 
cover associated with hardwood forest, but diversity 


392 


was lower relative to the conifer forest. More research 
is required to explain these differences. 

There are some limitations to the sampling method- 
ology used in this study. Taxonomic levels lower than 
the one used here to identify invertebrates would cer- 
tainly provide more detailed results though sampling 
efficiency would be reduced. Sampling was done 
around mid-day, which limits sampling to invertebrate 
groups that are active during the day (Eisenbeis and 
Wichard 1987). Results are limited to the invertebrates 
visually observed (i.e., >1 mm which may exclude 
arthropods such as small mites) found under boxes 
whereas smaller individuals and groups found in deep- 
er soil are under-represented. Another potential con- 
cern is that ants, due to their clumped distribution and 
significant effects on community structure, may obscure 
patterns among other invertebrate groups (Madden 
and Fox 1997). 

Diversity indices respond to both changes in species 
richness estimates and changes in species evenness. 
For example, the mesic lowland hardwood site record- 
ed few RTU but high total density. Among those com- 
mon taxa associated with the mesic lowland sites are 
springtails, earthworms, and isopods, which might be 
expected to be more abundant in moist soils. As a result, 
one problem with interpretation of diversity results is 
the possible relationship between measures of richness 
and diversity relative to measures of density. Both 
Shannon’s and Simpson’s indices are expected to be 
lowest when richness and evenness are low and to 
increase with richness and evenness. Shannon’s index 
is more sensitive to richness and is impacted by the 
inclusion of rare taxa, while Simpson’s index is more 
sensitive to evenness of the more common taxa. The 
result is usually that the two are generally correlated, 
although the relative ranking of sites may differ some- 
what. Among the five forest types reported in Table 2, 
the two hardwood sites had the highest numbers of 
individuals. Both Shannon’s and Simpson’s indices are 
reported as moderate for these sites with higher esti- 
mates for both in the upland sites. The two conifer- 
grass sites yielded the lowest number of individuals. 
The Shannon’s estimate is the lowest of the five sites 
while the Simpson’s is the highest. Between the other 
two sites, which recorded somewhat higher numbers 
of individuals, the Shannon’s estimates are the two 
highest while the Simpson’s estimates are the two low- 
est. The net effect is that across all five habitat types 
the correlation coefficient between the Shannon and 
Simpson estimates for the forest stands arranged along 
a moisture gradient had a negative slope, although not 
significant, contrary to expectations. However, both 
indices were negatively correlated with % herb cover 
and positively correlated with each other. Greater repli- 
cates are required in various forest stands along envi- 
ronmental gradients to account for forest characteris- 
tics that covary. 

Although increasing plant diversity significantly 
increases invertebrate diversity, local herbivore diversi- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


ty is also maintained by a diversity of parasites (e.g., 
flies and nematodes) and predators (Siemann et al. 
1998; Ferguson 2001). A community-level prediction 
is that invertebrate diversity increases with increasing 
plant species diversity, as many invertebrates forage 
on the leaves and litter of herbaceous plants (Symstad 
et al. 2000). However, if an entire functional group of 
plants, such as conifers, is absent from a habitat, then 
a landscape-level decrease in soil invertebrate diver- 
sity would occur, as many taxa are associated with the 
acidic soil and fungal hyphae characteristic of conif- 
erous forest floors (Lattin 1993; Butterfield 1999). 
Alternatively, diversity is hypothesized to increase and 
decrease across the two dimensions of productivity and 
disturbance, respectively (Kondoh 2001). Our results 
conform to this view, as conifer cover had the highest 
diversity at the late successional stage but the lowest 
measure of diversity at the early conifer-grass stage. 
In contrast, more constant mesic hardwood habitats 
had high productivity and low disturbance, and showed 
intermediate indices of invertebrate diversity. Appar- 
ently, the pattern of invertebrate community diversity 
varied among habitat types according to productivity 
and disturbance gradients. 

The decrease in soil invertebrate diversity with 
increasing grass cover may be due to the low nutri- 
tive value of grass and the microhabitat conditions of 
grassy areas. Low moisture, associated with dry grassy 
areas, has been found to have an adverse effect on soil 
invertebrate abundance due to a reduced oxygen level 
that degrades soil composition, increases erosion, and 
mobilizes carbohydrates and nutrients (Marra and 
Edmonds 1998). The majority of soil invertebrates are 
best suited to moderate moisture levels, as some spe- 
cies have little or no exoskeleton (e.g., earthworms) to 
protect against high and low soil moisture (Schaefer 
1995; Ferguson 2004). 

Soil invertebrate diversity was not significantly 
related to many of the forest characteristics. Still, the 
considerable diversity of invertebrates ensures that 
some species are adapted to many of the diverse con- 
ditions. For example, soil invertebrates did not show 
a relationship to the number of downed logs. This 
lack of a relationship with logs may have been relat- 
ed to the experimental technique of providing a simi- 
lar microhabitat using boxes. Many soil invertebrates 
are known to depend on cover provided by coarse 
woody debris, such as downed logs, due to their pro- 
vision of nutrients and protection from predators and 
stability of microclimatic conditions (Lattin 1993; 
Ferguson and Joly 2002). Slugs showed few relation- 
ships with forest characteristics and were found in a 
diversity of habitats. Slugs forage on larvae of bee- 
tles and flies and on cellulose and other plant poly- 
saccharides (Port and Port 1986). Earthworms bur- 
row in moist rich soil and feed on decaying organic 
matter from fallen leaves and vegetation (Edwards 
and Bohlen 1996). Earthworms were found to be 
positively associated with herbaceous cover, nega- 


2004 


tively associated with conifer cover, and positively 
associated with soil moisture. The major food items 
of centipedes are earthworms and small arthropods 
(Formanowicz and Bradley 1987). Although cen- 
tipedes have been found to prefer moist habitats 
(Corey and Stout 1992), such as under logs, no sig- 
nificant relationships were found with the forest 
characteristics measured in this study. 

Springtails are abundant in soil, and consume 
decomposing plant material and fungal hyphae (Hop- 
kin 1997). Their abundance was negatively associat- 
ed with litter cover, perhaps because it is living annu- 
al plants that provide the overwinter dead material 
that fungal hyphae provide as springtail food. The 
non-significant correlation (P = 0.06) of springtail 
abundance with herbaceous cover and the negative 
association with conifer composition contradicts 
other studies (Butterfield 1999; Paquin and Coderre 
1997). A possible explanation for these differences is 
the differing sampling methodologies used whereby 
surface-dwelling springtails were sampled here, in 
contrast to sub-surface sampling from other studies. 

Beetles showed little dependence on any of the 
measured forest characteristics in this study, in con- 
trast to another study that found habitat dependencies 
(Fournier and Loreau 2001). Beetles are adapted to a 
wide range of environments partly due to their 
exoskeleton enabling life in a variety of moisture 
conditions (Eisenbeis and Wichard 1987). Isopods 
occur in greater abundance under stones and in damp 
environments (Sutton 1980), which is confirmed in 
this study by the positive association with relative 
soil moisture. Also, isopods were positively associat- 
ed with herbaceous versus conifer cover, which is 
related to intolerance to acidic conditions of conifer 
soils and the greater food availability in deciduous 
and herbaceous forest cover (David et al. 2001). 
These findings differ with previous research that 
found isopod abundance unrelated to forest attributes 
(Bolger et al. 2000). 

Forest management, with the goal of preserving 
forest biodiversity that includes soil invertebrates, 
needs to consider the requirements of individual taxa 
in boreal ecosystems by providing varied landscapes 
within a mosaic of forest stands. For example, cli- 
mate change may profoundly influence boreal forest 
ecosystems and their management, via increased 
temperature and altered precipitation regimes (e.g., 
fires, etc.; Parker et al. 2000). In the boreal forest 
area studied here, conifer habitat had the highest 
diversity, although abundance and RTU richness 
were lower. However, our results need to be guarded- 
ly interpreted as they are based upon observations of 
a partly artificial system and our invertebrate diversi- 
ty results are not sufficiently comprehensive to draw 
management conclusions. Still, monitoring soil 
invertebrate diversity can provide a means to assess 
changes in forest environments with climate warm- 


FERGUSON and BERUBE: INVERTEBRATE DIVERSITY 393 


ing as well as forest management practices that 
include silvicultural interventions to maintain forest 
health. 

Acknowledgments 


Students in the “Fish and Wildlife Practice” 4" year 
course offered at Lakehead University participated in 
data analysis. Financial support was provided by post- 
doctoral (SHF) and postgraduate (DKAB) funding from 
NSERC (Natural Sciences and Engineering Research 
Council) and Bowater Pulp and Paper Inc., Thunder 
Bay Woodlands Division. 


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Simpson, E. H. 1949. Measurement of diversity. Nature 163: 
688. 

Sutton, S. L. 1980. Woodlice. Pergamon Press, Oxford. ISBN: 
0-080-25942-1. 

Symstad, A. J., E. Siemann, and J. Haarstad. 2000. An 
experimental test of the effect of plant functional group 
diversity on arthropod diversity. Oikos 89: 243-253. 

Tilman, D., and S. Pacala. 1993. The maintenance of species 
diversity in plant communities. Pages 13-25 in Species 
diversity in ecological communities. Edited by R. Ricklefs, 
and D. Schluter. University of Chicago Press, Chicago. 
ISBN: 0-226-71823-9. 

Wallwork, J. A. 1983. Oribatids in forest ecosystems. Annual 
Review of Entomology 2: 1016-1023. 


Received 16 April 2003 
Accepted 23 November 2004 


Observations of Habitat Use by Polar Bears, Ursus maritimus, in the 
Alaskan Beaufort, Chukchi, and Northern Bering Seas 


DONALD J. HANSEN 
U.S. Department of the Interior, Minerals Management Service, 949 E. 36'" Avenue, Anchorage, Alaska 99504 USA 


Hansen, Donald J. 2004. Observations of habitat use by Polar Bears, Ursus maritimus, in the Alaskan Beaufort, Chukchi, 
and northern Bering Seas. Canadian Field-Naturalist 118(3): 395-399. 


A total of 1 112 Polar Bears (Ursus maritimus) at 482 sightings were recorded during aerial surveys in the Beaufort, Chukchi, 
and northern Bering Seas conducted primarily during September and October from 1979-1999. Of these bears, 784 were 
observed offshore at 400 sightings. The surveys were conducted by the Naval Ocean Systems Center and Minerals Manage- 
ment Service; they were designed to monitor the fall Bowhead Whale (Balaena mysticetus) migration. Over the 20-year 
period, 1 096 620 kilometers of surveys were flown. The majority of the offshore Polar Bears, 595 bears at 290 sightings, 
and most of the kill sites and polar bear tracks were recorded in 80-100% ice cover. The number of bears per kilometer 
increased substantially in >24% ice cover, with the highest number observed in 80-100% ice cover. This habitat use proba- 
bly is related to the availability of seals, their primary prey. There were 328 bears (83 sightings) recorded on land, and most 
of them were associated with whale carcasses and bowhead whale subsistence harvest sites along the Alaskan Beaufort Sea coast. 


Key Words: Polar Bear, Ursus maritimus, habitat, Beaufort Sea, Chukchi Sea, Northern Bering Sea, Alaska. 


The preference of Polar Bears for extensive ice 
habitats is well documented (Stirling et al. 1993; Stir- 
ling and Derocher 1993; Amstrup 1995). Polar Bears 
need an ice platform to hunt successfully for seals, 
their primary prey (Stirling, Andriashek, and Calvert 
1993). The bears select ice habitats that increase ac- 
cessibility (availability) of seals. Ferguson et al. (2000) 
compared ice-habitat use by Polar Bears of the Cana- 
dian Arctic Archipelago and Baffin Bay. They found 
ice-habitat use to be similar for the two regions. How- 
ever, Baffin Bay bears had more limited access to 
Ringed Seals (Phoca hispida) because of the complete 
absence of ice cover during the summer season. The 
Canadian Arctic Archipelago had more consistent sea- 
sonal ice cover, which provided the bears with reli- 
able access to the seals. Although the Baffin Bay region 
had a much higher density of Ringed Seals (1.4-2.1 
seals/square kilometer: Finley et al. 1983; Stirling and 
Oritsland 1995) than the Arctic Archipelago region 
(0.28-0.97seals/square kilometer: Kingsley, Stirling, 
and Calvert 1985), both regions had very similar Polar 
Bear densities. Ferguson et al. (2000) concluded that 
Polar Bear abundance is not always proportional to prey 
density because of differences in Polar Bear access to 
seals (i.e., availability of prey). 

During September and October (open-water season), 
the Alaskan Beaufort Sea has great variability in ice 
coverage from year to year (Figures la and 1b). Like 
Baffin Bay, the Arctic coast of Alaska during light ice 
years is free of ice during the open-water season, when 
the pack ice is more than 100 miles from the coast (see 
Figure la). During the absence of ice in coastal Beau- 
fort Sea habitats, bears that occur along the coast either 
scavenge on animal carcasses that wash ashore, feed 


on the remains of whale carcasses at subsistence whale 
harvest sites, or fast until shorefast ice forms and seals 
become available (United States Department of the 
Interior, Fish and Wildlife Service 1995). 

During heavy ice years, the pack ice remains on 
the continental shelf; ice floes are common in coastal 
waters, and shorefast ice remains or forms early dur- 
ing the fall (Figure 1b). These varying ice conditions 
affected the movements and distribution of Polar Bears 
in the Beaufort and Chukchi seas (Amstrup 2000). 


Methods and Materials 

From 1979-1999, the Naval Ocean Systems Center 
under contract with the Bureau of Land Management 
(1979-1982) and Minerals Management Service (1982- 
1999) under the Bowhead Whale Aerial Survey Project 
has conducted aerial surveys in the Beaufort, Chukchi, 
and Bering seas (Ljungblad et al. 1986; Moore and 
Clarke 1992; Treacy 2000) to monitor the fall migration 
(late August through October) of endangered whales 
(Treacy 2000). Information on Polar Bears and sea-ice 
coverage was acquired mostly from these aerial surveys. 
Surveys were flown primarily at 1500 feet (458 meters), 
weather permitting, or at a minimum of 1000 feet 
(305 meters) to avoid disturbing marine mammals. The 
aircraft used was a de Havilland Twin Otter Series 300 
with bubble windows on both sides of the plane. Earlier 
flights (1979-1991) over the northern Bering, Chukchi, 
and western Beaufort seas used a Grumman Turbo 
Goose model G21G (Ljungblad et al. 1986; Moore 
and Clarke 1992). 

Information also was used from aerial whale sur- 
veys conducted during the spring and summer (April 
through August) in the northern Bering Sea as well 


395 


396 


Arctic North Slope of Alaska 


THE CANADIAN FIELD-NATURALIST 


Maximum Retreat of the Pack Ice on 2 October 1998 


Vol. 118 


“600 Miles 


Ficure la. Light Ice Year Information derived from the National Naval Ice Center (http://www.natice.noaa.gov/westarct1.htm). 


Maximum Retreat of the Pack Ice on 18 September 1983 


Beaufort Sea 


Arctic North Slope of Alaska 


Maximum Retreat of the Pack Ice on September 18, 1983 


ak. 


Ficure 1b. Heavy Ice Year information derived from the National Naval Ice Center (http://www.natice.noaa.gov/westarct1.htm). 


as in the Chukchi and Beaufort seas (Ljungblad et al. 
1986; Moore and Clarke 1992). Incidental sightings 
of Polar Bears and their behavior were recorded along 
north-south transects and off transects along the re- 


mainder of the flight tracks (Figure 2). Information 
was recorded on percent ice coverage within about 1- 
2 kilometers of the aircraft at each position update 
(once about every 5 minutes) and at each sighting loca- 


| 
1 


2004 


ioe. 


Chukchi Sea 


Alaska 


Bering Sea 
& 


HANSEN: HABITAT USE BY POLAR BEARS 


397 


Polar Bear Sightings 


Survey Track 


FiGure 2. Study are with survey tracks and Polar Bear sightings. 


tion. Position updates, sighting locations, and ice- 
coverage data were recorded on an onboard comput- 
er system connected to a Global Positioning System 
receiver in the aircraft. These records included infor- 
mation on numbers of Polar Bears per sighting and 
ice cover within about 1-2 kilometers of the aircraft. 
Polar Bear tracks and kill sites also were recorded dur- 
ing September and October from 1987 through 1999. 

Survey effort in the Beaufort Sea focused primarily 
along the coast, continental shelf, and shelf break, while 
surveys in the northern Bering and eastern Chukchi seas 
extended from the coast to the International Date Line 
(Figure 2). The Polar Bear sightings, kill sites, locations 
of bear tracks, and ice-coverage data were converted 
into database files and analyzed with Arcview Geo- 
graphic Information System Program 3.2a (devel- 
oped by ESRI in Redland, Calif.). Tables and graphs 
were created in Microsoft Word and Microsoft Excel 
(Windows 2000) to compare bear numbers and sight- 


ings with ice coverage (Table | and Figure 3) and kill 
sites and bear tracks with ice cover (Figures 4 and 5). 
The number of bear sightings and the survey effort 
(kilometers flown) were plotted by ice category to com- 
pare ice-cover use (Figure 2). This analysis is provid- 
ing useful information on habitat use by Polar Bears 
occurring in Alaskan waters. 


Results and Discussion 

Table 1 shows the number of Polar Bears and num- 
ber of sightings recorded offshore by ice cover and 
on the mainland or on barrier islands. Figure 4 shows 
the number of Polar Bear kill sites and ice coverage 
recorded at the kill locations. Figure 5 shows the num- 
bers of Polar Bear tracks recorded and ice coverage 
at their locations. 

Over the 20-year period and the 1 096 620 kilome- 
ters of surveys flown, 1112 Polar Bears at 482 sightings 
were recorded in the Beaufort, Chukchi, and Bering 


398 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE |. Number of Polar Bears and Number of Sightings (Recorded Per Year) by Ice Coverage and Number Recorded on 


Land. 

Year 80-100% Ice 50-79% 25-49% 
Ice Ice Ice 

1979 15 (8) 

1980 29 (2) ayy) iy, <6) 

1981 S005) Sii(Q) Dale) 

1982 16.3) 4 (4) ives) 

1983 165 365) 14 (11) Brn) 

1984 23)a- 1 CLS) (2) Sy) 

1985 5 (3) ihe (Cb) 

1986 9 (6) 3y2@) 

1987 4 (2) i eo Gh) 

1988 69 (43) 1G) PEO) 

1989 155) 0) a= 9A) One®) 

1990 52 (40) Perth) 

1991 DD ran 3) Aa (2) Gd) 

1992 LOS Gib) Sie G2) 

1993 Sh aC) 

1994 26°, (12) PO) 32) 

1995 Disi(eld) 

1996 9 (7) ta) 

1997 53 (8) 1) ED) 

1998 15 (6) 9 (4) 1b Gb) 

1999 2 (2) 1b) 

Totals: 5957 1(290) 91 (52) 45 (23) 


Polar Bear Sightings/Kilometer 


@ bear/km 


79-50% 49-25%  — 24-1% 0% 
Ranges of Ice Coverage 


100-80% 


Ficure 3. Polar Bear sightings per kilometer recorded from 
April 1979 through October 1999 in the Beaufort, 
Chukchi and Bering Seas. 


Ice Coverages at Kill Sites 


1 
{ 


77) 
o 
= 
2) 
S 
- 
) 
_ 
o 
Q 
= 
3 
rs 


100% 99-95% 94-90% 89-80% 79-70% 69-50% 49-40% 


Percent Ice Coverage 


39-1% 


FiGuRE 4. Number of kill sites and ice coveage recorded at 
each site. 


1-24% 
Ice No Ice 


Total Number of 
On Land Bears and Sightings 
51) 20 (10) 
(29) 
(20) 
1 (1) 22 (19) 
93 (49) 
3-7(2) et) S50 1@2) 
6 (4) 
(10) 
356) 
72 (46) 
(25) 
53 (41) 
(18) 
(48) 
Sp (@) 
(16) 
2 @) 
10 (8) 
LL @) 65 (11) 
217 (57) 247 (73) 
33 (19) 39 (25) 
328 (83) 1112 (482) 


67 (5) 


8 (5) 
S13) 
28 17) 25, (7) 


seas from 1979-1999 (Table 1). Of these bears, 784 
were observed offshore at 400 sightings. Figure 2 
shows the number of Polar Bears per kilometer of 
survey effort over ranges of ice coverage. The num- 
ber of bears per kilometer increased substantially in 
>24% ice cover, with the highest number observed in 
80-100% ice cover. These observations suggest that 
Polar Bears may use 80-100% ice cover more often 
than lower percentages of ice cover. This habitat use 
probably is related to the availability of seals, their pri- 
mary prey. Table | lists the number of bears and sight- 
ings seen offshore in other ice concentrations. Thirty- 
five kill sites and 769 Polar Bear track sightings were 
recorded, mostly in 80-100% ice cover (Figures 3 and 
4, respectively). Tracks with zero ice cover were record- 
ed on land or barrier islands. 

A total of 328 bears (at 83 sightings) were recorded 
on the mainland or on barrier islands (Table 1). Most 
of these bears were associated with whale carcasses 
and bowhead whale subsistence harvest sites along 
the Beaufort Sea coast. The large numbers of bears 
were recorded on land in 1992 (67 bears) and 1998 
(217 bears), when repeat sightings were made of sever- 
al bears feeding on whale carcasses along the Beau- 
fort Sea coast (Table 1). 

Studies of radio- and satellite-tagged Polar Bears 
have provided useful information on the movements 
and habitat use of individual Polar Bears (Arthur et 
al. 1996; Ferguson et al. 1998; Amstrup et al. 2001; 
Durner et al. 2004). These studies recorded Polar Bear 
locations and compared them with satellite-derived, 


2004 


Number of Polar Bear Tracks 


nm 
a 
ao 


nm 
o 


n 
x 
rs) 
£ 
(= 
° 
- 
o 
a 
= 
Ss 
z 


100 99-95 94-90 89-80 79-70 69-60 59-50 49-40 39-30 29-20 19-1% 0 
Percent Ice Coverage 


FiGurE 5. Number of Polar Bear tracks and ice coverage 
recorded at each site. 


digitized maps of ice types and ice concentrations in 


_ the general areas where the bears were located. These 


remote-sensing techniques provide general informa- 
tion on habitat use by Polar Bears throughout the 
year. Aerial surveys in this study recorded ice types 
and ice concentrations within 1-2 kilometers of the 
aircraft along the flight track and at the specific loca- 
tions where Polar Bears were observed. Both tech- 
niques record information on habitat use. Aerial sur- 
veys provide real-time information on habitat use at 
the actual locations where the bears were observed, 


_ while remote-sensing techniques provide valuable 


information on bear movements and general informa- 
tion on habitat use over time. 

Many aerial surveys conducted specifically for 
Polar Bears are flown at lower altitudes than the 1500 
foot surveys conducted in this study. Aerial surveys 
that are conducted at very low altitudes (200-300 feet) 
are very likely to disturb Polar Bears and other wild- 
life. Our surveys were conducted at 1500 feet to avoid 
disturbing marine mammals. Even at 1500 feet some 


_ of the bears observed reacted to the aircraft by run- 


ning away when the aircraft past overhead. Polar Bears 
are quite visible on the ice at 1500 feet because their 
tracks are very detectable. Other marine mammals of 
comparable size to Polar Bears, such as Beluga Whales, 
are easily detected at 1500 feet. 


_ Acknowledgments 


This study was funded by the U.S. Department of 
the Interior, Minerals Management Service, as part 
of its Alaska Environmental Studies Program. As the 
Principal Investigator, I would like to thank Stephen 
Treacy, Project Manager for the Minerals Manage- 
ment Service Bowhead Whale Aerial Survey Project, 
for providing the survey data on Polar Bears and ice 
coverage along the survey tracks from 1979 through 
2000, and for his comments on the draft manuscript. 
I also would like to thank Elinore M. Anker, who edited 
the manuscript. 


Literature Cited 
Amstrup, S. C. 1995. Movements, distribution, and population 
dynamics of Polar Bears in the Beaufort Sea. Ph.D. thesis. 


HANSEN: HABITAT USE BY POLAR BEARS 


399 


University of Alaska, Fairbanks, Fairbanks, Alaska. 299 
pages. 

Amstrup, S. C. 2000. Polar Bear. Pages 133-157 in Chapter 
7 in The natural history of an Arctic oil field development 
and the biota. Edited by J.C. Truett and S. R. Johnson. 
Academic Press, San Diego, California. 

Amstrup, S. C., G. M. Durner, T. L. McDonald, D. M. 
Mulcahy, and G. W. Garner. 2001. Comparing movement 
patterns of satellite-tagged male and female Polar Bears. 
Canadian Journal of Zoology 79: 2147-2158. 

Arthur, S. M., B. F. J. Manly, L. L. McDonald, and G. W. 
Garner. 1996. Assessing habitat selection when avail- 
ability changes. Ecology 77: 215-227. 

Durner, G. M., S. C. Amstrup, R. Neilson, and T. McDon- 
ald. 2004. The use of sea ice habitat by female Polar Bears 
in the Beaufort Sea. OCS Study MMS 2004-014. USDOL, 
MMS, Alaska OCS Region, Anchorage, Alaska. 41 pages. 

Ferguson, S. H., M. K. Taylor, and F. Messier. 2000. Influ- 
ence of sea ice dynamics on habitat selection by Polar 
Bears. Ecology 81: 761-772. 

Ferguson. S. H., M. K. Taylor, E. W. Born, and F. Messier. 
1998. Fractals, sea-ice landscape and special patterns of 
Polar Bears. Journal of Biogeography 25: 1081-1092. 

Finley, K. J., G. W. Miller, R. A. Davis, and W. R. Koski. 
1983. A distinctive large breeding population of Ringed 
Seals (Phoca hispida) inhabiting the Baffin Bay pack ice. 
Arctic 36: 162-173. 

Kingsley, M. C. S., I. Stirling, and W. Calvert. 1985. The 
distribution and abundance of seals in the Canadian high 
Arctic, 1980-1985. Canadian Journal of Fisheries and 
Aquatic Science 42: 1189-1210. 

Ljungblad, D. K., S. E. Moore, J. T. Clarke, and J. C. Ben- 
nett. 1986. Aerial surveys of endangered whales in the north- 
em Bering, eastern Chukchi, and Alaskan Beaufort Seas, 
1985: with a seven year review, 1979-85. OCS Study, MMS 
86-0002. USDOI, MMS, Alaska OCS Region, Anchorage, 
Alaska. 142 pages. 

Moore, S. E., and J. T. Clarke. 1992. Distribution, abundance, 
and behavior of endangered whales in the Alaskan Chukchi 
and western Beaufort Seas, 1991: with a review 1982-91. 
OCS Study, MMS 92-0029. USDOI, MMS, Alaska OCS 
Region, Anchorage, Alaska. 126 pages. 

Stirling, I., and A. E. Derocher. 1993. Possible impacts of 
climatic warming on Polar Bears. Arctic 46: 240-245. 
Stirling, I., D. Andriashek, and W. Calvert. 1993. Habitat 
preferences of Polar Bears in western Canadian Arctic in 

late winter and spring. Polar Record 29(168): 13-24. 

Stirling, I., and N. A. Oritsland. 1995. Relationships between 
estimates of Ringed Seal (Phoca hispida) and Polar Bear 
(Ursus maritimus) populations in the Canadian Arctic. 
Canadian Journal of Fisheries and Aquatic Sciences 52: 
2594-2612. 

Treacy, S. D. 2000. Aerial surveys of endangered whales in 
the Beaufort Sea fall 1998, and 1999. OCS Study, MMS 
2000-066. USDOI, MMS, Alaska OCS Region, Anchor- 
age, Alaska. 135 pages. 

USDOI, Fish and Wildlife Service. 1995. Habitat conser- 
vation strategy for Polar Bears in Alaska. USDOI, Fish and 
Wildlife Service, Alaska Region, Anchorage, Alaska. 119 
pages. 


Received 23 April 2003 
Accepted 16 August 2004 


New Records for the Arctic Shrew, Sorex arcticus and the Newly 
Recognized Maritime Shrew, Sorex maritimensis 


Nett D. Perry!, DONALD T. STEWART?, ELIZABETH M. MADDEN®, and THOMAS J. MAIER* 


'Department of Wildlife and Fisheries Science, Texas A&M University, 210 Nagle Hall, College Station, Texas, 77843-2258 
USA. Corresponding author. 

“Department of Biology, Acadia University, Wolfville, Nova Scotia, B4P 2R6 Canada 

‘U.S. Fish and Wildlife Service, Medicine Lake National Wildlife Refuge, 223 North Shore Road, Medicine Lake, Montana 
59247 USA 

USDA Forest Service, Northeastern Research Station, University of Massachusetts, Amherst, Massachusetts 01003-9285 
USA 


Perry, Neil D., Donald T. Stewart, Elizabeth M. Madden, and Thomas J. Maier. 2004. New records for the Arctic Shrew, Sorex 
arcticus, and the newly recognized Maritime Shrew, Sorex maritimensis. Canadian Field-Naturalist 118(3): 400-404. 


We report the first record for the Arctic Shrew (Sorex arcticus) in the state of Montana, USA. We also report range extensions 
for the closely related Maritime Shrew (Sorex maritimensis) in New Brunswick and Nova Scotia, Canada. These collections 
augment our limited knowledge of the ranges and habitat associations of these rarely collected shrews, and highlight the 
need for a careful assessment of the status of S. maritimensis in Canada. 


Key Words: Arctic Shrew, Sorex arcticus, Maritime Shrew, Sorex maritimensis, range, state record, Montana, New Brunswick, 


Nova Scotia, Quebec. 


Approximately 38 species of shrews (genus Sorex) 
are currently recognized in North America (Hall 1981; 
Jones et al. 1986; George 1988; Wolsan and Hutterer 
1998; Fumagalli et al. 1999). Most North American 
shrews belong to the subgenus Otisorex, which is pri- 
marily restricted to this continent. Until recently, the 
only two recognized members of the subgenus Sorex 
found in North America were the Tundra Shrew (Sorex 
tundrensis) and the Arctic Shrew (Sorex arcticus) (van 
Zyll de Jong 1983a), the latter species documented 
throughout much of the boreal forest region of North 
America. Recent molecular work, however, has support- 
ed the recognition of the Maritime Shrew (Sorex mari- 
timensis, previously S$. arcticus maritimensis) as a 
distinct species (Stewart et al. 2002). The range of the 
Maritime Shrew is limited to the eastern portion of 
New Brunswick and Nova Scotia. 

Sorex arcticus and S. maritimensis are infrequently 
observed or collected. Both species exhibit a preference 
for grass-sedge meadows and wetland edges (Wrigley 
et al. 1979; van Zyll de Jong 1983b; Kirkland and 
Schmidt 1996). Although there are limited studies which 
include estimates of density (Buckner 1966), both spe- 
cies appear to exist at lower population densities than 
other common mammal species of the boreal region 
(e.g., Masked Shrew [Sorex cinereus] and Meadow 
Vole [Microtus pennsylvanicus]), although they may 
be locally common in appropriate habitat (Wrigley et 
al. 1979). Sorex maritimensis, because of its limited 
range and restriction to areas of grass-sedges, is con- 
sidered rare (van Zyll de Jong 1983b). This may be a 
result of competitive exclusion by similar sized Smoky 
Shrews (Sorex fumeus) which exhibit a preference for 
less mesic, wooded habitats (van Zyll de Jong 1983a). 


Herein we report the first record of S. arcticus in 
Montana and range extensions of S. maritimensis in 
New Brunswick and Nova Scotia, and provide addi- 
tional data on habitat associations in these locations. 
We also report a collection of S. arcticus in Sept Hes, 
Quebec, one of only six records within the province 
(Peterson 1966; van Zyll de Jong 1983b). In the follow- 
ing sections, we detail trapping methods and results 
for each collection. 


Methods 

Montana, USA. Between 22 and 27 July 2001 we 
conducted small mammal baseline surveys on wet 
meadow habitats at Medicine Lake National Wildlife 
Refuge, Sheridan County, northeast Montana (48°30'N, 
104°20'W; Figure 1). The refuge is located in the gla- 
cially influenced prairie pothole region, noted for its 
gentle rolling plains with occasional shallow depres- 
sions—host to vast wetlands and seasonally flooded 
meadows. 

We set two 150 m transects, each with 10 trap sta- 
tions set 15 m apart for five consecutive nights. Each 
station included three different Victor® snap-traps: 
one mouse trap, one museum special trap, and one rat 
trap (300 trap nights). Nearby pitfall traps consisted of 
two 5-m fences (aluminum flashing) with 5-L paint 
buckets (dry, not baited) at either end (40 trap nights; 
each night a bucket was open was considered a trap 
night). The transects were located in the Lake Creek 
flood plain, an area seasonally inundated with water 
during spring run-off and early summer rains (Stuart 
and Kantrud 1971). The plant community was herba- 
ceous, dominated by sedges (Carex spp.), grasses (Agro- 
pyron spp. and Spartina spp.), and rushes (Juncus 


400 


2004 


PERRY, STEWART, MADDEN, and MAIER: ARCTIC AND MARINE SHREWS 


40] 


FicurE 1. Suggested range for Sorex arcticus. Includes extensions presented herein and ranges suggested by Peterson 
(1966) and van Zyll de Jong (1983b). Insets: A + Indicates location of Montana S. arcticus collections, Medicine 
Lake National Wildlife Refuge. @ Indicates location of nearest known record, Lostwood National Wildlife Refuge, 
North Dakota. B. o Indicates the location of the Quebec S. arcticus collection, other points (@) indicate location of 
historical collections in Quebec. 


spp.), with a variety of wetland forbs interspersed. Iden- 
tical survey efforts were conducted in two additional 
habitat types on the refuge: native prairie and planted 
perennial grasslands. Total combined effort for all three 
habitat types was 900 snap trap nights and 120 pitfall 
nights. 

Quebec, Canada. One hundred pitfall traps (800 trap 
nights) were set at each of two sites, 17-21 July 1990 
(Figure 1). Sites were 8 km north and 8 km east of Sépt 
Iles, Quebec (50°12'N, 66°23'W) in coniferous wood- 
land near the edge of a marsh and in an old grassy field 
next to a road, respectively. The coniferous woodland/ 
marsh was characterized by White Spruce (Picea 
glauca), Balsam Fir (Abies balsamea) and sedges 
(Carex spp.). 

New Brunswick, Canada. A total of 175 non-baited 
pitfall traps (525 trap nights) were set 3 km southeast 
of St. George, near L’Etete, New Brunswick (45°8'N, 


66°50'W), 3-7 August 1990 (Figure 2). Traps were set 
in thickets of predominantly alder (Alnus sp.) with some 
mixed conifer growth and some grasses. A small brook 
(0.5 m wide) transected the site, maintaining relatively 
moist soils. 

Nova Scotia, Canada. [Method information is not 
available] 


Results 

Montana, USA. Six S. arcticus, the first confirma- 
tion of this species in Montana (Foresman 2001), were 
collected in wet meadows at Medicine Lake National 
Wildlife Refuge (Figure 1). The nearest known previous 
collection was at Lostwood National Wildlife Refuge, 
Burke and Mountrail counties, North Dakota, approx- 
imately 190 km to the east (R. Murphy, personal com- 
munication). 


402 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


200 Kilometers 
_ 


FiGureE 2. Suggested range of Sorex maritimensis, including collections described herein. A indicates the location of the St. 
George collection, New Brunswick. B indicates location of the Belle Isle collection, Nova Scotia. Historic locations, 
indicated by solid dots, were taken from van Zyll de Jong (1983b). We found no extralimital collections since that 


publication. 


These specimens were captured using Victor® Mu- 
seum Specials (n = 5) and Victor® rat traps (n = 1) 
baited with a mixture of peanut butter, oatmeal, flour, 
and black sunflower seeds. No S. arcticus were col- 
lected in either pitfall traps or smaller Victor® mouse- 
traps. Although identical survey efforts were conduct- 
ed in native prairie and planted perennial grasslands, 
S. arcticus were collected only in wet meadow habi- 
tats. Other species collected from wet meadow sites 
included: Meadow Vole (n = 19), Deer Mouse (Pero- 
myscus maniculatus, n = 1), and Masked Shrew (n = 8). 

All specimens were confirmed as S. arcticus; how- 
ever, they were collected on the periphery of the S. a. 
arcticus and S. a. laricorum ranges, and identification 
to subspecies is difficult. These specimens are cata- 
logued at the Philip L. Wright Zoological Museum at 
the University of Montana, Missoula (catalog numbers 
UMZM 18554 — 18559). 

Quebec, Canada. Two Arctic Shrews (S. a. arcticus; 
Royal Ontario Museum [ROM] catalogue numbers 
110254 and 110255) were collected by D.T-.S. near 
Sept Iles, Quebec (50°12'N, 66°23'W), 17-21 July 1990 
(Figure 1). This is the sixth reported collection of S. 
arcticus in this province (Peterson 1966; van Zyll de 
Jong 1983b). Both specimens were collected from the 
grassy site east of Sept Iles. Other species collected 
during this effort were Masked Shrew (n = 7), Pygmy 
Shrew (Sorex hoyi; n = 1), and Meadow Vole (n = 1). 


This record is as far north along the immediate 
coast of the St. Lawrence River as S. arcticus have 
previously been reported. The last specimen in this 
general area, near Moisie River, just north of Sept 
Iles, was trapped in 1937 (van Zyll de Jong 1983b). 
Though little trapping has been conducted in this 
region, van Zyll de Jong (1983b) speculates that the 
species is distributed further north in Quebec and 
Labrador throughout the boreal forest, which includes 
extensive marshy habitats. 

New Brunswick, Canada. Three Maritime Shrews 
(originally identified as S. a. maritimensis; ROM cata- 
logue numbers 110314, 110315, 110331) were col- 
lected 3 km southeast of St. George, near L’Etete, New 
Brunswick (45°8'N, 66°50'W), 3-7 August 1990 (Fig- 
ure 2). This collection extends the known range of S. 
maritimensis ca. 100 km south. Other species collected 
from this effort were Masked Shrew (n = 21) and 
Northern Short-tailed Shrew (Blarina brevicauda; 
e—1))' 

Nova Scotia, Canada. On 4 October 1992, a single 
specimen of Sorex maritimensis (originally identified 
as §. a. maritimensis) was collected at Belle Isle, 
Nova Scotia (Tom Herman, personal communication). 
The specimen, collected in a marshy area dominated 
by the sedge Scirpus cyperinus, represents a provin- 
cial range extension of this species by ca.100 km 
(Figure 2). 


2004 


Discussion 

Sorex arcticus. The range of the Arctic Shrew is 
strongly associated with the boreal forest region of 
North America. Southward range expansions of four 
other boreal species, Masked Shrew, Meadow Vole, 
Meadow Jumping Mouse (Zapus hudsonius), and the 
Least Weasel (Mustela nivalis), have been correlated 
with contemporary cool, mesic climate patterns in the 
Great Plains region (Frey 1992). Jannett and Huber 
(1994) speculate that a recent southward extension of 
S. arcticus in Minnesota is associated with these cool- 
ing climate patterns. Indeed, this southwestward exten- 
sion of S. arcticus into Montana could be correlated 
with this phenomenon. 

Nevertheless, few small mammal surveys have been 
conducted in northeastern Montana (D. Flath, person- 
al communication); it is possible this has prevented 
earlier detection in the state. Similar wet meadow 
habitat exists approximately 40 km to the south as 
the Big Muddy Creek feeds into the Missouri River. 
Thus, it is plausible that the range of S. arcticus ex- 
tends further into Montana. 

All Montana S. arcticus specimens were sexually 
inactive young-of-year. Clough (1963) observed that 
over-wintering S. arcticus captured between February 
and July were reproductively active. Also, the char- 
acteristic tri-colored pelage for adults of this species 
was indistinct in these specimens, further supporting 
our designation of these shrews as young-of-year, 
products of early season breeding by the previous 
year’s cohort (Clough 1963; Baird et al. 1983). These 
specimens were collected in two groups of three, 
each group approximately 3 km apart and separated 
by a small perennial stream. Given their inactive re- 
productive status and that individuals from each group 
were collected no greater than 15 m apart, group mem- 
bers might have been litter-mates. As such, a viable 
population of S. arcticus likely exists at Medicine 
Lake National Wildlife Refuge. 

Sorex maritimensis. The Maritime Shrew was pre- 
viously considered restricted to the north and east of 
the St. John River system, with the nearest previous 
collection for this species near Saint John, New Bruns- 
wick (Peterson 1966; van Zyll de Jong 1983b). The 
collection of S$. maritimensis near L’ Etete, New Bruns- 
wick, brings this species within 30 km of the Maine 
border. Maine and New Brunswick are separated in 
this area by the St. Croix River which could be a bar- 
rier to shrew dispersal. There is suitable habitat for S. 
maritimensis on the U.S. side of the border and it is 
possible they will be found there; however, to date there 
is no record of S. maritimensis in Maine (J. Albright, 
R. Boone, and L. Master, personal communications). 
S. maritimensis is currently recognized as one of only 
four mammals endemic to Canada (the others are the 
Varying Lemmings /Dicrostonyx hudsonius D. richard- 
soni], the Gaspé Shrew [Sorex gaspensis], and the 
Vancouver Marmot [Marmota vancouverensis]). 


PERRY, STEWART, MADDEN, and MAIER: ARCTIC AND MARINE SHREWS 


403 


The Nova Scotia and New Brunswick collections of 
the newly recognized S. maritimensis imply a greater 
range, extending inland and likely including the entire- 
ty of mainland Nova Scotia. Stewart et al. (2002) 
speculate that recent glacial encroachment (ca. 20 000 
y) may have isolated this species on the coastal flood 
plain of Nova Scotia. These recent collections may 
either provide evidence that S. maritimensis is re- 
colonizing former range, following the reestablish- 
ment of the boreal forest after the last ice age, or may 
reflect the paucity of efforts to collect S. maritimensis 
within suitable habitats in these provinces. These col- 
lections also support a hypothesis that this species is 
limited to moist grasslands and bogs associated with 
the boreal forest, competitively excluded from habitats 
occupied by the Smokey Shrew, a closely related wood- 
land associate. This limited distribution and restriction 
to fragile wet meadow habitats suggests this species 
may warrant conservation concern. 

These collections offer some data regarding the range 
and niche characteristics of these shrews. Given that 
S. arcticus and S$. maritimensis have been the focus 
of very few research efforts, more research targeting 
specific life history traits are necessary to better under- 
stand the habitat associations and range restrictions 
of these shrew species. 


Acknowledgments 

We particularly thank Tom Herman for providing 
data on his specimen of S. maritimensis collected in 
Nova Scotia and Kerry Foresman for facilitating the 
processing and accession of Montana specimens into 
the University of Montana mammal collection. We also 
thank the staff at Medicine Lake National Wildlife 
Refuge for funding and support of small mammal col- 
lections on the refuge. We thank D. Flath and R. Mur- 
phy for insight into the status of S. arcticus in Mon- 
tana and North Dakota. DTS thanks S. Hindocha and 
H. Russell for assistance in the field; and J. Albright, 
R. Boone, and L. Master for comments on the status 
of S. maritimensis in Maine. The research program of 
DTS is supported by an NSERC Discovery grant. 


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Buckner, C. H. 1966. Populations and ecological relation- 
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Clough, G. C. 1963. Biology of the Arctic Shrew, Sorex 
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Foresman, K. R. 2001. The wild mammals of Montana. 
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Frey, J. K. 1992. Response of a mammalian faunal element 
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George, S. B. 1988. Systematics, historical biogeography, and 
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Hall, E. R. 1981. The mammals of North America. Volume 1. 
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Jones, J. K., Jr., D. C. Carter, H. H. Genoways, R. S. 
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Kirkland, G. L. Jr., and D. F. Schmidt. 1996. Sorex arcticus. 
Mammalian Species (524): 1-5. 

Peterson, R. L. 1966. The mammals of eastern Canada. 
Oxford University Press, Toronto. 465 pages. 

Stewart, D. T., N. D. Perry, and L. Fumagalli. 2002. The 
Maritime Shrew, Sorex maritimensis (Soricidae: Insecti- 


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vora): a newly recognized, Canadian endemic. Canadian 
Journal of Zoology 80: 94-99. 

Stuart, R. E., and H. A. Kantrud. 1971. Classification of 
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van Zyll de Jong, C. G. 1983a. A morphometric analysis of 
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special consideration of the taxonomic status of S. a. 
maritimensis. Le Naturaliste Canadien 110: 373-378. 

van Zyll de Jong, C. G. 1983b. Handbook of Canadian mam- 
mals. Volume |. Marsupials and insectivores. National 
Museums of Canada, Ottawa. 210 pages. 

Wolsan, M., and R. Hutterer. 1998. A list of the living spe- 
cies of shrews. Appendix in Evolution of shrews. Edited 
by J. M. Wojcik and M. Wolsan. Mammal Research Insti- 
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Habitat, abundance, and distribution of six species of shrews 
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Received 12 May 2003 
Accepted 21 December 2004 


Lichen Zonation on Coastal Rocks in Gwaii Haanas National Park 
Reserve, Haida Gwaii (Queen Charlotte Islands), British Columbia 


IRWIN M. Bropo! and Norm A. SLOAN2 


‘Canadian Museum of Nature, P. O. Box 3443 Station D, Ottawa, Ontario K1P 6P4 Canada 
> Parks Canada, Gwaii Haanas National Park Reserve and Haida Heritage Site, P. O. Box 37, Queen Charlotte City, British 
Columbia VOT 1SO Canada 


Brodo, Irwin M., and Norm A. Sloan. 2004. Lichen zonation on coastal rocks in Gwaii Haanas National Park Reserve. Haida 
Gwaii (Queen Charlotte Islands), British Columbia. Canadian Field-Naturalist 118(3): 405-424. 


The occurrence of 43 marine lichen species on intertidal rocky shores of southern Haida Gwaii (Queen Charlotte Islands), 
British Columbia is described and related particularly to elevation on the shore (duration of seawater immersion) and 
exposure to waves. In the area of Gwaii Haanas National Park Reserve and Haida Heritage Site on Moresby Island, rock- 
dwelling marine lichens are distributed in zones much as they are elsewhere in the world, although some species found 
abundantly only in Haida Gwaii give the local shores a unique appearance. In common with other areas, there is a 
conspicuous black band of Verrucaria species (in this case, nine species plus other black lichens) at the upper edge of the 
intertidal zone. A conspicuous white band of Coccotrema maritimum above the black band is a unique feature of this flora. 
The unusually large percentage of endemic, near-endemic or disjunct lichen species and their phytogeography suggest that at 
least the headland rocks along the west coast were refugia during the last glacial maximum. Verrucaria striatula and V. 
sandstedei are reported for the first time from British Columbia. 


Key Words: Maritime lichens, zonation, Haida Gwaii, Pacific Northwest, British Columbia. 


Lichens form a dominant, yet often overlooked, 
component of the maritime vegetation along rocky sea- 
shores. They create distinct bands along such shores 
from polar to tropical seas with maximum abundance, 
and the most studies, in north-temperate regions (Flet- 
cher 1980; Little and Kitching 1996). The species res- 
ponsible for these bands in the northeast Pacific (1.e., 
the Pacific Northwest of North America; see the cover 
photo, and Plate 64 in Brodo et. al. 2001: 77), however, 
have rarely been investigated. 

The shoreline lichen vegetation of Gwaii Haanas 
National Park Reserve and Haida Heritage Site can be 
understood in the context of the total lichen flora of 
Haida Gwaii, British Columbia. Five field trips by the 
senior author to Haida Gwaii since 1967 have revealed 
a diverse lichen flora of about 580 species with more 
than 25 new to science (Brodo 1995). Many of the spe- 
cies are rare worldwide, and more than a dozen are en- 
demic to Haida Gwaii or are known only from a few 
nearby localities in southeast Alaska and Vancouver 
Island. Further, many species are disjuncts, having their 
nearest additional locality thousands of kilometres dis- 
tant, often in western Europe, Asia or the southern 
hemisphere (Brodo 1992). 

With the establishment of Gwaii Haanas as a Na- 
tional Park Reserve, there is a need to develop a base- 
line inventory of lichen species for various uses, for 
example, to facilitate the conservation of lichen bio- 
diversity (Hunter and Webb 2002), to track human and 
natural perturbations, to assist in nature interpretation, 
and to aid studies of shoreline ecology. Lichens are 
well-known indicators of environmental health in use 
world-wide (Nimis et al. 2002). Regionally, they have 


been used for air quality monitoring in the Tongass 
National Forest in southeast Alaska (Geiser et el. 1994*). 
In addition, lichen studies have been made to evaluate 
the effects of oil spills and post-spill cleaning on marine 
ecosystems (Lallemant and Van Haluwyn 1981; Flet- 
cher and Crump 2002). A knowledge of maritime lich- 
ens is clearly relevant to the prospect of oil and gas 
development in the Hecate Strait separating Haida 
Gwaii from the British Columbia mainland (Anony- 
mous 2002*). 

It is important to know if the shoreline lichens of 
Haida Gwaii differ from those found elsewhere on the 
west coast of North America. The only other detailed 
study of Pacific Northwest marine lichen zonation was 
done by Ryan (1988a, 1988b) in Washington State, al- 
though some observations on shoreline lichens from 
southern British Columbia were presented by Noble 
(1982). 

We report here on the distribution of lichen species 
in the upper rocky intertidal zone landward into the salt 
spray zone in the Gwaii Haanas area. Because splash 
patterns and exposure affect establishment and growth 
of shoreline lichens, we examined rocky shores with a 
range of exposures to wave action from “protected” to 
“very exposed” to reveal species occurrence patterns. 
Finally, we compared the rock-dwelling marine lichen 
flora and zonation of southern Haida Gwaii with the 
flora and patterns seen in other regions. 


Study Area 

Haida Gwaii, known also as the Queen Charlotte 
Islands, is an archipelago of about 138 islands in the 
hypermaritime zone of the Pacific Northwest, lying 


405 


406 


about 80 km off the mainland British Columbia coast 
between Vancouver Island and southeast Alaska (Fig- 
ure |). Its geography, climate and vegetation were des- 
cribed in some detail in Brodo (1995). 

The two largest land masses of Haida Gwaii are 
Graham Island to the north and Moresby Island to the 
south. The lower elevations of Graham Island (the 
eastern two-thirds) and Moresby Island (the northern 
third) have been disturbed by intensive logging activ- 
ities, but the southern two-thirds of Moresby Island 
were set aside in 1988 as a National Park Reserve 
called Gwaii Haanas lying roughly between 52° and 
53°N longitude (Figure 2) covering ca. 1470 km? of 
land and ca. 1700 km of shoreline (Sloan and Bartier 
2000). The lichen studies described here mainly centre 
on the extreme southern end of the Park Reserve, at 
the tip of Moresby Island and on Kunghit Island (Fig- 
ures 2 and 3). 


Methods 

To characterize our sample locations, we used Gwaii 
Haanas’ shoreline classification system (Harper et al. 
1994*). This is in the park’s geographic information sys- 
tem (GIS) and is based largely upon substrate type, 
texture and exposure to wave action. This system was 
first developed in Gwaii Haanas and has since become 
the standard physical shoreline classification system of 
British Columbia (Howes et al. 1994*) and Washington 
State [http://www2.wadnr.gov/nearshore/index.asp]. 
The system has a biological zoning component in 
which visible lichens are coarsely grouped into a 
“Verrucaria band” (Searing et al. 1995). The British 
Columbia scheme has been used in broad-scale region- 
al studies of intertidal species diversity (Zacharias and 
Roff 2001). 

In Harper et al. (1994*), lichens are mentioned as 
forming conspicuous shoreline bands. There are refer- 
ences to the “Verrucaria zone” and the “white lichen 
zone,” but the diversity of species is overlooked. For 
example, it is not made clear that the Verrucaria zone 
consists of nine black Verrucaria species and other 
dark lichens (Brodo and Santesson 1997), and that the 
white zone is caused by the dominance of Coccotrema 
maritimum, a species described from Haida Gwaii, 
whose closest relatives are in Asia and the southern 
hemisphere. It is rare in the Pacific Northwest outside 
Haida Gwaii and adjacent coastal regions (Brodo 
1973). Other lichens of intertidal rocks also proved to 
be new to science and generally restricted to Haida 
Gwaii and nearby coastal localities. These include 
Caloplaca litoricola, Fuscidea thomsonii, Verrucaria 
schofieldii, V. epimaura, and Porina pacifica (Brodo 
1984: Brodo and Wirth 1998; Brodo and Santesson 
1997; Brodo 2004). Several other novelties remain to 
be formally described. 

Fourteen locations within Gwaii Haanas and one on 
Limestone Island were sampled (Figures 2, 3). At each 
location, one to three transects were laid out for a total 
of 18 transects. A 12 m tape measure was deployed 


THE CANADIAN FIELD-NATURALIST 


Vol. 117 


conforming closely to the rock surface and perpendi- 
cular to the high tide line. Transects usually began at 
the upper limit of the rockweed (Fucus gardneri) and 
barnacle (Balanus and Semibalanus spp.) zone (the 
“FB zone,” approximating the mean high water level 
[MHW)]) and ran landward to the point at which terres- 
trial vascular plants or mosses began. This distance 
was divided into eight or nine equal sections to give 
approximately nine to ten quadrat points. Because ex- 
posed, wave-splashed beaches and shallow slopes pro- 
duced longer transects than those on protected, rela- 
tively steep shores, the increments of height above the 
FB zone from quadrat to quadrat were not uniform, 
nor were the distances between quadrats. 

At each quadrat point, a 10 x 10 cm grid on trans- 
parent plastic with a hundred 1-cm? squares, was cen- 
tered on the transect tape. All lichen species under the 
grid were recorded together with their coverage in 
percent (each square representing 1% coverage). For 
smooth rocks, the coverage readings were fairly accu- 
rate, but with increasing surface roughness, the esti- 
mates were correspondingly approximate. After the 
quadrats were sampled and the lichen data recorded, 
the vertical height of each quadrat above the starting 
point in the FB zone was recorded. 


Results 

The 18 transects were sorted and analyzed accord- 
ing to degree of shoreline exposure from Protected to 
Very Exposed, based on Harper et al. (1994*), as listed 
in Table 1. Two locations had multiple transects, there 
were no transects for localities 6 or 10, and transect 
11 had incomplete data and was excluded. Limestone 
transects are indicated (L) and all other transects were 
over mixed volcanic and fine-grained sedimentary rocks 
characteristic of the region’s shoreline (Sutherland 
Brown and Yorath 1989). 

The transects in Gwaii Haanas yielded 43 lichen 
species. Sixty-seven rock-dwelling species occurring 
on maritime rocks are currently known from Haida 
Gwaii as a whole. An annotated list of all shoreline 
species in Haida Gwaii is presented as an Appendix. 
This updates the list in Sloan and Bartier (2000; Ap- 
pendix B, Part 2) compiled by the senior author based 
on specimens at the National Herbarium of Canada 
(CANL), Ottawa. The zones indicated in Sloan and 
Bartier (2000) were assigned tentatively, based on 
label data, and have been corrected in this study. Two 
of the lichens from our transects (Verrucaria striatula 
and V. sandstedei) are new lichens for Haida Gwaii 
(and British Columbia), and ten others were not includ- 
ed in the Sloan and Bartier (2000) list because they 
were not regarded to be maritime. These are: Adelole- 
cia kolaensis, Gyalecta jenensis, Opegrapha gyro- 
carpa, Parmelia saxatilis, Placopsis lambii, Porpidia 
contraponenda, P. speirea and P. thomsonii. Many rela- 
tively common maritime and marine lichens found 
elsewhere in Haida Gwaii were not seen in the Gwaii 
Haanas transects. These include Caloplaca inconspecta, 


2003 BRODO AND SLOAN: LICHEN ZONATION ON COASTAL ROCKS 407 


Langara 1327 'W 


Island Dixon Entrance 


Canada 
Masset 
e 
f , British 


A Columbia 
| + ak 


Graham 
Island 


Hecate 
Queen 
Charlotte : 
Citys. Skidegate Strait 
® Sandspit 


Louise 
Island 
Moresby 


Island 


GD 


Gwaii Haanas Burnaby 


A Island 
National Park Reserve 


and Haida Heritage Site / 
proposed National Marine 
Conservation Area 


SGaang 
Gwaii Kunghit 
Island 
Queen 
100 Km Cape Charlotte 
St. James Sound 


FicurE 1. Location of Gwaii Haanas within Haida Gwaii (Queen Charlotte Islands) including some of the larger islands. 


408 


132°W 


Louise 
Island 


va 


Moresby 
Island 


Gwaii Haanas 
National Park Reserve 


and Haida Heritage Site 
and proposed 
National Marine 
Conservation Area 


THE CANADIAN FIELD-NATURALIST 


Vol. 117 


131°W 


Ron 


18 Limestone 
Island 


Lyell Hecate 


Island eunah 
Strait 


Burnaby 
Island 


Inset 2 


17 
Kunghit Treat Bay 


Island 


GOON 


50 Kilometres 


132°W 


131°W 


FIGURE 2. Map of the Gwaii Haanas area showing Limestone Island (location 18) to the north and Treat Bay (location 17) 


to the south of the insets (see Figure 3 for insets). 


Catillaria calybeia, Rhizocarpon hensseniae and Rino- 
dina gennari. At least one lichen (Porina pacifica) was 
found growing abundantly just outside the quadrats 
on a protected shore. 

The frequency of occurrence (%) of each species 
in each transect is listed according to shore exposure 
class in Table 2. Verrucaria maura occurred in all tran- 
sects on all shores. Among the eight most abundant 
lichens (frequencies exceeding 50% in all transects), 


four are endemic or nearly endemic to Haida Gwaii: 
Caloplaca litoricola, Coccotrema maritimum, Verru- 
caria epimaura and V. schofieldii; and one is endemic 
to the coastal Pacific Northwest (Herteliana alasken- 
sis). The semi-exposed shores had the most species and 
the most exposed shores yielded the fewest species. 
The results of the transect observations, expressed as 
metres above the FB (Fucus/barnacle) zone, are given 
in Tables 3 and 4 organized according to the degree of 


2003 BRODO AND SLOAN: LICHEN ZONATION ON COASTAL ROCKS 


; Ee 
; F bY e TS phelican Foint Tnset 


pre 
wie, eas 
Skincuttle han 
i 4 ; ee : Hecate 
: \ # &. a Ka 


Strait 


ns Gamenten 8 7 - Ingraham Point 
Bay ray 


¢ 


5 Kilometres 


Houston Stewart 
Channel 


Islands 


4 Kilometres 
SSS 


Ficure 3. Inset maps from Figure 2 showing lichen sampling locations 1-16 within Gwaii Haanas (see Table 1). 


409 


410 


THE CANADIAN FIELD-NATURALIST 


Vol. 117 


TABLE 1. Locations of shore lichen transects in the Gwaii Haanas area grouped according to the shore exposure classification 


of Harper et al. (1994). 


Exposure Class Location Name 


Transect No.* 


Latitude (°N) Longitude (°W) 


Protected Carpenter Bay 9 131.1660 52.2312 
SGaang Gwaii 13 13022511 S2a1015 
Limestone Island 18(L>) 131.6170 52.9086 
Semi-protected Ellen Island 1 131.0918 52.1544 
Orion Point 5) 131.0231 52.1229 
Semi-exposed Gordon Islands 2-1, 2-2, 2-3 131.1443 52.0967 
Orion Point + 131.0217 52.1241 
Pelican Point 14 (L’) 131.2615 52.3472 
Pelican Point 15 131.2618 52.3458 
Houston-Stewart Channel 16-1, 16-2 131.1261 21258 
Treat Bay ity) 131.0080 52.0659 
Exposed Ingraham Point 7 131.0325 32.232) 
Ingraham Point 8 131.0331 52.2319 
Very exposed Gordon Islands 3 131.1493 52.0965 
SGaang Gwaii 1 131.2330 52.1021 


4 Two locations (2 and 16) had multiple transects, locations 6 and 10 had no transects, the transect at location 11 was in- 


complete and excluded. 
>L = limestone rock 


exposure of each shore. These data are graphed Figures 
in 4 and 5, with the ordination based on average mini- 
mum values (the base of the thick black line). Because 
the values for all shore exposures except “Very Ex- 
posed” appear to largely overlap with respect to the 
height above the FB zone, the data are pooled in Table 
3 and Figure 4; Table 4 and Figure 5 shows the data 
only for Very Exposed shores. The data for Quadrat 
Position (Table 5), being relative, are not greatly affect- 
ed by shore exposure, so all shores are considered 
together. The results of all transects with 8-10 quadrats 
per transect are shown in Figure 6. (Transects having 
only 6 or 7 quadrats cannot be compared with those 
having 8-10 transects.) 

Translating the continuum of species position into a 
zonal scheme with names that correspond with previ- 
ous North American or European studies proved to be 
challenging. The graphs presented in Figures 4 and 5 
based on metres above the upper limit of the FB zone 
(Tables 3 and 4) were compared to Figure 6 showing 
the species according to their quadrat placement along 
the transects (Table 5) with the aim of finding natural 
breaks in the continua. This was only partially suc- 
cessful. Thus, in Table 6, which presents a summary of 
the zones, their position and their dominant lichens, a 
few species are listed twice, in different zonal cate- 
gories, according to height above the Fucus-barnacle 
limit versus their relative quadrat position. Our zonal 
terminology basically follows the British System, es- 
pecially Fletcher (1973a, 1973b) who, in agreement 
with Lewis (1961), argued that the zones are best 
described and defined on biological criteria, not on 
physical (i.e., tidal) criteria. Lewis (1961) recommend- 
ed adopting “Littoral Fringe” in place of “Supralit- 
toral Fringe” used by Stephenson and Stephenson 
(1949) because it better describes the community that 
inhabits the upper fringe of the intertidal zone, washed 


by most tides in relatively protected shorelines. We 
replace Mesic, Submesic and Xeric subdivisions of the 
Supralittoral Zone with the more easily understood 
terms Lower, Middle and Upper, respectively. In addi- 
tion, we find that the term “xeric,” even taken in its 
relative sense, is not an accurate description of a shore- 
line habitat. Ryan (1988b) used a numbering system 
for his zones, but they basically followed the British 
scheme following the distribution of shoreline organ- 
isms. O’Clair et al. (1996) used a simplified system 
based on wave exposure, and this was adopted by Sloan 
and Bartier (2000). Some zonal synonymy, discussed 
in detail by Ryan (1988b), is given in the first column 
of Table 6. 

In the Gwaii Haanas area, Verrucaria mucosa and 
Pyrenocollema halodytes are the most abundant spe- 
cies at the upper edge of the intertidal zone; i.e., the 
lowest point on our transects, where rockweed or bar- 
nacles occur (Lower Littoral Fringe; quadrat 1). They 
grow there together with some of the smaller and rarer 
species of Verrucaria such as V. sandstedei and V. stri- 
atula. These are generally too small, thin, and scattered 
to create a black zone and, therefore, are hard to find. 
The black zone begins with the appearance of the 
blacker, thicker species of Verrucaria: V. erichsenii, 
V. maura, V. epimaura and V. schofieldii, all of which 
can have broad coverage (Table 7) in the Upper Lit- 
toral Fringe (quadrats 2-4). Although V. maura con- 
tinues to be important well up on the shoreline, the 
community changes its character and color with the 
appearance of the gray-brown, gray, and orange species 
such as Herteliana alaskensis, Arthonia phaeobaea, 
and the near-endemic but abundant Haida Gwaii lichen, 
Caloplaca litoricola, in the Lower Supralittoral Zone 
(quadrats 5-7). Also in this zone is Adelolecia kolaensis 
and the cyanophilic lichen Spilonema revertens. Above 
this is a poorly defined orange zone (Middle Supra- 


EE qC“eer_ 


2003 


BRODO AND SLOAN: LICHEN ZONATION ON COASTAL ROCKS 


41] 


TABLE 2. Percentage occurrence of shoreline lichen species at locations according to shore exposure classification of Harper et al. 
(1994) in the Gwaii Haanas area. P = protected; SP = semi-protected; SE = semi-exposed; E = exposed; VE = very exposed. 


% occurrence according to shore exposure class % 


Lichen species P (3) SP (2) 
Verrucaria maura 100 100 
Spilonema revertens 67 100 
Caloplaca litoricola 33 100 
Coccotrema maritimum 67 50 
Verrucaria epimaura 0 100 
Herteliana alaskensis 67 100 
Pyrenocollema halodytes 67 0 
Verrucaria schofieldii 67 100 
Arthonia phaeobaea 33 50 
Verrucaria amphibia 100 0 
Lecidella stigmatea 67 100 
Rhizocarpon geminatum 33 50 
Xanthoria candelaria 0 50 
Amandinea coniops 0 100 
Physcia caesia 33 50 
Porpidia speirea 0 

Verrucaria erichsenii 0 

Verrucaria mucosa 67 

Caloplaca flavogranulosa 33 

Collema fecundum 33 


Fuscidea thomsonii 
Adelolecia kolaensis 
Verrucaria degelii 
Verrucaria sandstedei 
Lecanora muralis 
Ochrolechia subplicans 
Parmelia saxatilis 
Placopsis lambii 
Porpidia thomsonii 
Rhizocarpon haidensis ined. 
Aspicilia caesiocinerea 
Caloplaca citrina 
Caloplaca rosei 
Caloplaca verruculifera 
Candelariella sp. 
Fuscopannaria maritima 
Gyalecta jenensis 
Lecanora sp. #2 
Lecanora sp. #1 
Lecanora straminea 
Opegrapha gyrocarpa 
Opegrapha sp. 

Porina pacifica 

Porpidia carlottiana 
Porpidia contraponenda 
Rhizocarpon hochstetteri s. str 
Verrucaria sp. #5 
Verrucaria striatula 


i>) 
Sees 
Nn 


Nn 


Nn Nn 
SESS nO SIS OSS 2 OS 2 SEO OES TOO So. OrO rr Orr Oo Orr 


WwW 
SrOL ISTO Dru iOrO LOO OS OOS Os OOS O20 FSGS 
nn nN 


ies) 
nn 


Oo 


— 
\o 
i) 


Total lichen species 


littoral Zone; quadrats 7-8) with Caloplaca flavogran- 
ulosa, C. rosei (both west coast endemics) and the 
widespread Xanthoria candelaria, as well as the in- 
conspicuous crust, Rhizocarpon “haidensis.” The 
Upper Supralittoral Zone (quadrats 8 — 9) almost at 
the upper edge of the beach is well marked by the dom- 
inance of the pinkish white lichen, Coccotrema mari- 


(number of transects) occurrence in 


SE (9) E (2) VE (2) all transects 
100 100 100 100 
56 100 50 67 
56 100 50 61 
67 100 0 61 
67 50 100 61 
33 100 50 56 
56 50 100 56 
56 0 0 50 
33 100 50 44 
33 50 0 39 
0 0 100 33 
22, 0 50 28 
22 0 100 28 
22 0 0 22 
22 0 0 22 
44 0 0 22 
33 0 50 22 
yp) 0 0 22 
22, 0 0 17 
11 50 0 17 
22 50 0 17 
0 50 0 17 
33 0 0 17 
33 0 0 17 
0 0 50 11 
11 50 0 11 
11 50 0 11 
22 0 0 11 
11 50 0 11 
22 0 0 11 
0) 0 0) 6 
11 0 0 6 
0 0 0 6 
0 0 50 6 
0) 0 0) 6 
11 0 0 6 
11 0 0 6 
0 0 0 6 
0 0 0 6 
0 0 50 6 
0 0 0 6 
11 0 0 6 
0 0 0 6 
0 0 0 6 
11 0 0 6 
0 0 0 6 
11 0 0 6 
0 0 50 6 
33 15 15 


timum, which forms a conspicuous white band on the 
coastal rocks of Haida Gwaii, especially on the Pacific 
side, although it is less common outside of the archi- 
pelago. Accompanying the Coccotrema are such wide- 
spread lichens as Placopsis lambii, various species of 
Rhizocarpon, and rarer west coast species such as Och- 
rolechia subplicans. Higher on the shore and influ- 


412 


THE CANADIAN FIELD-NATURALIST 


Vol. 117 


TABLE 3. Summary data for Protected to Exposed shores, metres above Fucus/barnacle zone (16 quadrats). 


Species # Extreme 
quadrants minimum 
Adelolecia kolaensis 3 135 
Arthonia phaeobaea 7 0.55 
Aspicilia caesiocinerea 1 2.05 
Caloplaca citrina 1 1.05 
Caloplaca flavogranulosa 3 1.24 
Caloplaca litoricola 10 0.55 
Caloplaca rosei 1 2.05 
Candelariella sp. 1 2.65, 
Coccotrema maritimum 11 1.10 
Collema fecundum 3} 0.80 
Fuscidea thomsonii 3 2.20 
Fuscopannaria maritima 1 2.40 
Gyalecta jenensis 1 0.90 
Herteliana (Bacidia) alaskensis 9 0.60 
Lecanora muralis 2 2.65 
Lecanora sp. no.1 1 0.55 
Lecanora sp. no.2 1 0.90 
Lecidella stigmatea 4 0.60 
Ochrolechia subplicans 2 1.80 
Opegrapha gyrocarpa 1 1.83 
Opegrapha sp. 1 1.80 
Parmelia saxatilis 2 2.00 
Physcia caesia 4 1.00 
Placopsis lambii p) 1.85 
Porina pacifica if 2.70 
Porpidia carlottiana 1 1.50 
Porpidia contraponenda 1 1.60 
Porpidia speirea + 1.10 
Porpidia thomsonii 2 2.00 
Pyrenocollema halodytes 8 -0.20 
Rhizocarpon geminatum 4 1.70 
Rhizocarpon haidensis y) E25) 
Rhizocarpon hochstetteri s. str 1 1.30 
Spilonema revertens 11 0.60 
Verrucaria amphibia 7 0.20 
Verrucaria degelii 3 0.30 
Verrucaria epimaura 9 0.25 
Verrucaria erichsenii 3 0.00 
Verrucaria maura 16 -0.20 
Verrucaria mucosa 4 -0.20 
Verrucaria sandstedei 3 0.00 
Verrucaria schofieldii 9 -0.15 
Verrucaria sp. #5 1 0.90 
Xanthoria candelaria 3 1.40 


enced only by intermittent salt spray, is the Terrestrial 
Zone (quadrats 9-10), characterized by widespread non- 
maritime but salt-tolerant species such as Parmelia 
saxatilis as well a Haida Gwaii endemic, Fuscidea 
thomsonii, and a few lichens restricted to the Pacific 
Northwest, such as Fuscopannaria maritima and Por- 
pidia carlottiana. The bird rock community (ornitho- 
coprophilous lichens), including Lecanora straminea, 
L. muralis, and Caloplaca verruculifera, also occurs 
in this zone. 

In comparing the graph showing zonation on Pro- 
tected to Exposed shores (Figure 4) with the graph deal- 
ing with Very Exposed shores (Figure 5), it is imme- 
diately apparent that on very exposed shorelines, most 


Average Average Extreme 
minimum (m) maximum (m) maximum (m) 
1.43 2.03 BES 
1.04 1.34 2.40 
2.05 3.15 3.15 
1.05 1.05 1.05 
1.85 1.85 2.20 
1.07 1.43 Balls 
2.05 2.65 2.65 
2.65 2.65 2.65 
2.05 239) 3.60 
1.40 1.50 1.90 
2.73 D213. 3.60 
2.40 2.40 2.40 
0.90 0.90 0.90 
1.08 1.20 2.45 
2.65 2.65 2.65 
0.55 0.55 0.55 
0.90 0.90 0.90 
25 1.30 2.05 
2.10 2.40 2.40 
1.83 1.83 1.83 
1.80 2.30 2.30 
2.20 2.20 2.40 
1.11 1.99 Sul 
1.93 Pil} 2.40 
2.70 3.60 3.60 
1.50 1.50 1.50 
1.60 1.60 1.60 
2.28 2.44 3.60 
2.05 2.05 2.10 
0.00 0.00 0.35 
2S 2.61 Bulld 
2.08 DSS 2.90 
1.30 1.30 1.30 
1.65 2.21 3.60 
0.44 0.61 0.69 
0.32 0.32 0.35 
0.45 0.81 DDS 
0.35 0.87 0.90 
0.45 1.93 3.60 
-0.05 0.08 0.20 
0.20 0.33 0.35 
0.62 0.78 1.50 
0.90 1.05 1.05 
1.97 233 3.65 


of the lichen species are much higher on the shore, the 
zones themselves are much broader (reflecting the 
height of wave splash), and not all species are in the 
same relative quadrat position. For example, the cyano- 
bacterial lichen, Spilonema revertens, which mainly 
occurs in the central section of the transects (Middle 
Suppralittoral) on Protected to Exposed shores, is found 
at the top of the transects (Terrestrial) on Very Exposed 
shores. Verrucaria maura begins to invade the rock 
over a metre above the Fucus/barnacle limit, and other 
shoreline lichens are proportionally higher on the 
shore. The five species at the top of the transects are 
terrestrial, but salt-tolerant, lichens, not really mari- 
time. Also unique to the Very Exposed shores are the 


2003 


BRODO AND SLOAN: LICHEN ZONATION ON COASTAL ROCKS 


413 


TABLE 4. Summary data for Very Exposed shores, metres above Fucus/barnacle zone (2 transects). 


Species # Extreme 
quadrants minimum 
Arthonia phaeobaea | 2.05 
Caloplaca litoricola | 2.05 
Caloplaca verruculifera I 3.65 
Herteliana alaskensis I 3.65 
Lecanora muralis | 6.70 
Lecanora straminea | 3.65 
Lecidella stigmatea 2, Poet fe) 
Pyrenocollema halodytes 2 0.00 
Rhizocarpon geminatum 1 6.70 
Spilonema revertens | 5.70 
Verrucaria epimaura 2 1-75 
Verrucaria erichsenii ] 0.70 
Verrucaria maura 2 1.10 
Verrucaria schofieldii I 1.10 
Verrucaria striatula | 0.00 
Xanthoria candelaria 2 3.65 


large areas of uncolonized rock (bare of lichens as 
well as barnacles or marine algae). 

The relative zonal position of some lichen species 
on protected shores differs when comparing relative 
quadrat position and actual distance above the FB zone 
(compare Figures 4 and 6). This may be due to the 
ability of these species to occupy broader zones than 
others, coupled with the method of averaging mini- 
mum and maximum values to draw the charts. The 
species included in more than one zonal category are 
marked with an asterisk (*) or exclamation (!) in 
Table 6. 

Species richness (diversity) appears to be greatest 
on Semi-exposed shores (33 species) but, since this 
category of exposure has the largest number of tran- 
sects (9), the significance of this number is difficult to 
evaluate. If one combines the lists in Table 2 for Pro- 
tected and Semi-protected shores (5 transects) and 
those of Exposed and Very Exposed shores (4 tran- 
sects), the numbers of species are closer: P+SP: 29 
species; SE: 33 species; E+VE: 23 species. Interest- 
ingly, the Very Exposed shores, with only 16 species, 
include 3 species found in no other exposure type, 
Verrucaria striatula, Caloplaca verruculifera and 
Lecanora straminea. 

One indication of species importance along the 
rocky shores is the ability of each species to form large 
patches, 1.e., with high coverage values. Table 7 sum- 
marizes the coverage classes of all the species that 
appeared in quadrats in this study. Some species, and 
even species groups, are clearly able to cover larger 
areas than others. Almost all the black Verrucaria spe- 
cies, for example, are able to cover over 50% of the 
100 cm? quadrats, which is why they form a conspic- 
uous black belt on the shore. Similarly, Coccotrema 
maritimum forms large patches as does Caloplaca 
litoricola and Ochrolechia subplicans. Some species 
never dominate their zones, however, always occur- 
ring in small patches even if they are frequently en- 


Extreme 
maximum (m) 


Average 
minimum (m) 


Average 
maximum (m) 


2.05 2.05 2.05 
2.05 3.65 3.65 
3.65 3.65 3.65 
3.65 3.65 3.65 
6.70 6.70 6.70 
3.65 3.65 3.65 
4.73 4.83 6.70 
0.00 0.00 0.00 
6.70 6.70 6.70 
5.70 6.70 6.70 
2:23 2.38 2.70 
0.70 1.70 1.70 
1.40 4.83 6.70 
1.10 1.10 1.10 
0.00 0.00 0.00 
5.18 5.18 6.70 


countered. Examples are Spilonema revertens, Pyreno- 
collema halodytes, Herteliana alaskensis and Arthonia 
phaeobaea. 

Rock type generally has a major influence on spe- 
cies composition in lichen communities, but some spe- 
cies are more tolerant of varying rock type than others. 
Verrucaria mucosa was especially abundant on lime- 
stone although present on siliceous rock as well. Some 
lichens found only, or most commonly, on the lime- 
stone transects were Verrucaria amphibia, V. species 
number 5, Caloplaca citrina and Gyalecta jenensis. 
Pyrenocollema halodytes occurs on limestone rocks as 
well as on barnacles (both rich in calcium carbonate). 
On the other hand, many common lichens; e.g., Cocco- 
trema maritimum, Caloplaca litoricola, Verrucaria 
epimaura, Herteliana alaskensis and Collema fecun- 
dum, were found only on siliceous rock. Species appar- 
ently oblivious to rock chemistry include Verrucaria 
maura and V. schofieldii. 


Discussion 

Marine and maritime lichens distribute themselves 
on rocky shorelines in response to many factors, with 
the dominant influences being wave exposure and, 
particularly, immersion duration (Southward 1958; 
Fletcher 1973a; Taylor 1982). Fletcher (1973a) said 
that lichens such as Verrucaria mucosa and V. striatula, 
which grow among the barnacles in the upper inter- 
tidal zone, require 52% submergence per year, the 
highest amount for any lichen. There is experimental 
evidence (Ramkaer 1977) that spore germination and 
hyphal growth at young stages of development are 
influenced by salt concentrations, and that the ability 
of different lichens to become established in shoreline 
habitats is positively correlated with their position on 
shoreline rocks. Those lowest on the shore (longest 
immersion duration) are the species that have best 
germination and hyphal growth under moderate or 
even high concentrations of seawater. Mature thalli 


414 


THE CANADIAN FIELD-NATURALIST 


Table 5. Summary data for relative quadrat position for all transects with 8-10 quadrats (15 transects). 


Species # Extreme 
quadrants minimum 
Adelolecia kolaensis 3 4.00 
Amandinea coniops 4 6.00 
Arthonia phaeobaea 7 4.00 
Aspicilia caesiocinerea 1 6.00 
Caloplaca flavogranulosa 3 6.00 
Caloplaca litoricola 10 4.00 
Caloplaca rosei 1 7.00 
Caloplaca verruculifera 1 9.00 
Candelariella sp. 1 7.00 
Coccotrema maritimum 9 6.00 
Collema fecundum 3 5.00 
Fuscidea thomsonii 3 9.00 
Fuscopannaria maritima 1 9.00 
Herteliana alaskensis 10 3.00 
Lecanora muralis 2 7.00 
Lecanora straminea 1 9.00 
Lecanora sp. no.1 1 4.00 
Lecanora sp. no.2 1 3.00 
Lecidella stigmatea 6 4.00 
Ochrolechia subplicans 2 7.00 
Opegrapha gyrocarpa 1 8.00 
Parmelia saxatilis 2 6.00 
Physcia caesia 3 6.00 
Placopsis lambii 2 8.00 
Porina pacifica 1 7.00 
Porpidia carlottiana 1 9.00 
Porpidia speirea 3 6.00 
Porpidia thomsonii 2 8.00 
Pyrenocollema halodytes 9 1.00 
Rhizocarpon geminatum 5 5.00 
Rhizocarpon haidensis 2 7.00 
Rhizocarpon hochstetteri s. str 1 8.00 
Spilonema revertens 11 5.00 
Verrucaria amphibia =) 1.00 
Verrucaria degelii 2 1.00 
Verrucaria epimaura 10 2.00 
Verrucaria erichsenii 4 1.00 
Verrucaria maura 15 1.00 
Verrucaria mucosa 2 1.00 
Verrucaria sandstedei 3 1.00 
Verrucaria schofieldii 8 1.00 
Verrucaria striatula 1 1.00 
Xanthoria candelaria 5 6.00 


might be affected by osmotic problems, acidity (sea 
water has a relatively high pH), and other seawater 
minerals. Other factors that influence zonation include 
degree of wetness and seepage of acidic fresh water 
from the land above the shore (Fletcher 1973b; Ryan 
1988b) and excess heating or freezing (Southward 
1958). Verrucaria maura occurs far up on exposed 
shores in depressions or shaded nooks indicating a 
requirement for moisture. This was noted by European 
workers, including Fletcher (1973b), who nevertheless 
regards V. maura as one of the more drought-tolerant 
of the marine Verrucaria species. Fletcher (1973a) 
found that V. maura actually has no requirement for 
submergence in sea water at all. It can survive with 


Vol. 117 
Average Average Extreme 
minimum (m) maximum (m) maximum (m) 
5.67 7.00 8.00 
6.50 6.50 7.00 
6.00 6.57 9.00 
6.00 8.00 8.00 
7.33 N33 8.00 
4.80 6.30 9.00 
7.00 7.00 7.00 
9.00 9.00 9.00 
7.00 7.00 7.00 
8.00 9.00 10.00 
6.00 6.67 7.00 
9.33 9.33 10.00 
9.00 9.00 9.00 
4.80 5.30 9.00 
7.50 7.50 8.00 
9.00 9.00 9.00 
4.00 4.00 4.00 
3.00 3.00 3.00 
6.17 6.67 8.00 
8.00 9.00 9.00 
8.00 8.00 8.00 
8.50 9.00 9.00 
6.00 7.00 8.00 
8.00 8.50 9.00 
7.00 8.00 8.00 
9.00 9.00 9.00 
8.00 8.67 10.00 
8.00 8.00 8.00 
1.00 1.00 1.00 
7.80 8.00 9.00 
7.00 9.50 10.00 
8.00 8.00 8.00 
6.27 7.73 10.00 
2.40 2.60 4.00 
1.00 1.00 1.00 
2.90 3.90 5.00 
DIDS 4.50 7.00 
2.47 UAL 10.00 
1.00 1.50 2.00 
1.00 1.67 3.00 
325 4.00 7.00 
1.00 1.00 1.00 
7.20 7.60 8.00 


only sea spray. Competition with algae and grazing 
animals (such as limpets and snails) is regarded to be 
a significant factor by some observers (Southward 
1958; Ryan 1988b) and minimal by others (Fletcher 
1973a). We did not see much evidence of grazing on 
the lichens of Gwaii Haanas. 

As noted by Fletcher (1980), the lichen bands can 
be characterized by life form as well as species. In the 
Littoral Fringe, all lichens are anatomically unstratified 
(i.e., without a differentiated cortex, algal layer and 
medulla) and almost all open by an ostiole rather than 
having a discoid apothecium. An exception in Gwaii 
Haanas is Herteliana alaskensis, a lichen with bia- 
torine apothecia found in the Lower Supralittoral Zone 


a 


2003 BRODO AND SLOAN: LICHEN ZONATION ON COASTAL ROCKS 415 


TABLE 6. Zones formed by lichens on maritime rocks in Gwaii Hanaas. The names of equivalent zones in O’ Clair et al. (1996) 
and Ryan (1988b) are also given. For species listed twice under “Characteristic lichens,” an asterisk (*) indicates the lichen’s 
zonal position according to its occurrence in the quadrat series (see Figure 6); an exclamation (!) indicates its zonal position 
according to its height above the Fucus-barnacle (FB) zone (see Figure 4). P to E = protected to exposed shores; VE = very 


exposed shores. 


Quad series m above 
Zone Type Quadrat FB zone 
Terrestrial (Halophilic) 9-10 2.3-3.6 
O’Clair: upper salt spray 
Ryan: Zones 2A- 2B 
Upper Supralittoral Zone 8-9 2.1-2.6 
O’Clair: mid salt spray 
Ryan: Zone 2C-2D 
Middle Supralittoral Zone 7-8 1.4-2.3 
O’Clair: low salt spray 
Ryan: Zone 2D 
Lower Supralittoral Zone 5-7 0.8-1.5 
O’ Clair: splash zone 
Ryan: Zone 2E 
Littoral Fringe, Upper 2-4 0.3-0.8 
O’Clair : high intertidal (p.p.) 
Ryan: Zone 3-2E 
Littoral Fringe, Lower 1-2 -0.3-0.3 


O’Clair: high intertidal (p.p.) 
Ryan: Zone 4A-3 


occasionally in the company of black species of Ver- 
rucaria. Lichens with apothecia, also having somewhat 
stratified crustose thalli, first appear in the Lower Su- 
pralittoral Zone. Higher still, in the Middle and Upper 
Supralittoral Zones, the stratified foliose lichens appear. 
The biological reasons for this phenomenon are fairly 
easy to surmise. Lichens that are submerged frequently 
have no need for a loosely organized medulla with air 


Characteristic lichens 
PtoE VE 


Parmelia saxatilis 
Fuscidea thomsonii 
Fuscopannaria maritima 
Rhizocarpon geminatum! 
Lecanora muralis! 
Porina pacifica 
Caloplaca verruculifera* 
Lecanora straminea* 
Porpidea speirea! 


Rhizocarpon geminatum 
Lecanora muralis 
Spilonema revertens 


Coccotrema maratimum 
Placopsis lambii 
Ochrolechia subplicans 
Xanthoria candelaria! 
Porpidia thomsonii 
Aspicilia caesiocinerea 
Rhizocarpon “haidensis”! 
Caloplaca rosei! 
Porpidea speirea* 
Rhizocarpon geminatum* 


Lecidella stigmatea 
Xanthoria candelaria 


Caloplaca flavogranulosa 
Caloplaca rosei* 
Rhizocarpon “haidensis” * 
Lecanora muralis* 
Xanthoria candelaria* 
Adelolecia kolaensis! 
Spilonema revertens! 


Caloplaca verruculifera 
Herteliana alaskensis 
Lecanora straminea 


Herteliana alaskensis 
Caloplaca litoricola 
Arthonia phaeobaea 
Physcia caesia 
Collema fecundum 
Adelolecia kolaensis* 
Spilonema revertens* 


Caloplaca litoricola 
Arthonia phaeobaea 
Verrucaria maura 
V.epimaura 


Verrucaria maura 
Verrucaria epimaura 
Verrucaria erichsenii 
Verrucaria schofieldii 
Verrucaria amphibia 


Verrucaria erichseniti 
V. schofieldii 


Pyrenocollema halodytes 
Verrucaria striatula 
Verrucaria mucosa 
Verrucaria sandstedei 


Pyrenocollema halodytes 
Verrucaria striatula 


spaces for gas exchange, and their spores are better 
distributed by oozing out of an ostiole than being shot 
into the air as would occur from an apothecial disk. 
Stratified apothecial lichens are adapted for a basically 
terrestrial life style. 

The shift in relative position in the zonation by 
Spilonema revertens on Very Exposed (where it is 
Terrestrial) as compared to less exposed shorelines 


Vol. 117 


Metres above Fucus/barnacles 


THE CANADIAN FIELD-NATURALIST 


416 


4.00 

3.50 - 

3.00 

2.50 

2.00 

1.50 

1.00 

0.50 

0.00 

-0.50 

2 2 O@ Sf & © oN DS 3 V © O © Ww @ Oo HS So © © @ oe & dS @ a > & > © © & / Oo & 
os os e) os ee © is & SES Oe” ss CLE SX o* SELES ES é es yy Sos eS 
LE COE EFC FEF BOE GF SEF OB EE ESE SF EEF EB E EP EE EES ES oF oF F EEF ESE 
@ as fs a” -® eo” SS @ 2S ee & S s Ra eS S oe @ SEOe PF oe & & 5 & Pes < Ss & & > we @ & os & eS 
PS eS SF FS SS SF PD GS FP OS SF SF So Pie Fe S ee & & P° QPS LY EF oF 
F : Oe 7 0 or 2 
oo “<. RS ws Rs Fo oe ae es Fe & 3 & Fee ss ror & QP x eS RO is) Ko s e  ¢ x ES eg Z S 
SF WANA RO w CN SC SS Se ie oe 
Ke ak es) 0 & ee PF ¥ of 
SS es Co we < ge 
x < « 
e € 


The thick line is the range of average values. The thin lines represent the total range (extreme maximum and 


FicureE 4. Zonation of maritime lichens on Protected to Exposed shores, ordered by metres above the Fucus-barnacle zone. 
minimum) values. 


2003 BRODO AND SLOAN: LICHEN ZONATION ON COASTAL ROCKS 417 


ecies etres above Fucus/barnacles 
mn Sp Metres above Fucus/barnacl 
“% 
Ge) 
% 
@ 
y ° A I) 7) rN on fo) Ni 00 
°% ° ° r=) ° ° ° ° ° ro) 
Pe ro) ro) ro) } ro) rs) ro) } } 


FiGuRE 5. Zonation of maritime lichens on Very Exposed shores, ordered by metres above the Fucus-barnacle zone. The 
thick line is the range of average values. The thin lines represent the total range (extreme maximum and minimum) 
values. 


418 THE CANADIAN FIELD-NATURALIST Vola? 


Species 4 Quadrant 
%, 
%e 
ke es y ° Nn h a oo S =a 
j=) S S 
beh ro) ros) ro) ) ) 8 S 


FiGcuRE 6. Zonation of maritime lichens on Protected to Very Exposed shores, ordered by relative quadrat position. The 
thick line is the range of average values. The thin lines represent the total range (extreme maximum and minimum) 
values. 


2003 


BRODO AND SLOAN: LICHEN ZONATION ON COASTAL ROCKS 


419 


TABLE 7. The number of quadrats sampled in which lichen species are represented, according to percentage coverage category. 


Coverage category (%) 


Lichen species 1-5 6-10 11-20 


Adelolecia kolaensis 4 | 2 

Amandinea coniops 3 

Arthonia phaeobaea 5 | | 

Aspicilia caesiocinerea | ] 

Caloplaca citrina 

Caloplaca flavogranulosa 1 

Caloplaca litoricola 6 4 3 

Caloplaca rosei 2 

Caloplaca verruculifera 1 

Candelariella sp. | 

Coccotrema maritimum 1 4 2 

Collema fecundum 

Fuscidea thomsonii 3 

Fuscopannaria maritima 1 

Gyalecta jenensis 1 

Herteliana alaskensis 7 4 3 

Lecanora muralis 2 

Lecanora straminea 1 

Lecanora sp. #1 

Lecanora sp. #2 

Lecidella stigmatea 4 

Ochrolechia subplicans 1 

Opegrapha gyrocarpa 

Opegrapha sp. 

Parmelia saxatilis 

Physcia caesia 1 

Placopsis lambii 2 1 

Porina pacifica 

Porpidia carlottiana 

Porpidia contraponenda 

Porpidia speirea 4 

Porpidia thomsonii 1 

Pyrenocollema halodytes 10 
3 
2 


— 
Sas a 


— 
i) 


BRNNRK eK 


— 


Rhizocarpon geminatum 
Rhizocarpon haidensis ined. 
Rhizocarpon hochstetteri s. str 1 
Spilonema revertens 17 
Verrucaria amphibia 2 
Verrucaria degelii 

Verrucaria epimaura 2 
Verrucaria erichsenti 1 
Verrucaria maura 11 
Verrucaria mucosa 

Verrucaria sandstedei 

Verrucaria schofieldii Z 2 2 
Verrucaria striatula 

Verrucaria sp. #5 

Xanthoria candelaria + 


i 
[id ) 


SN Oe 
aN 


— 
me 


(where it is Middle Supralittoral) may be due, in part, 
to the species’ sensitivity to excessive inundation by 
tidal splash. More likely, it is due to its inability to 
physically withstand the pounding of waves on highly 
exposed headland rocks. Spilonema is a microfruticose 
lichen and is attached to the rock relatively loosely 
compared to its crustose neighbours. A similar shift 
in the distribution of the fruticose Lichina pygmaea 
occurs on European shores (Fletcher 1973a). Spilo- 


21-30 


31-40 41-50 51-60 61-70 71-80 81-90 91-100 


w 
See nNWe 
oo 
tS 
— 
iT) 
NCO 


nema revertens is not a particularly maritime lichen, 
however, being frequently found in inland areas (Brodo 
et al. 2001). 

The species diversity on protected and semiprotect- 
ed Gwaii Haanas shores is not significantly lower than 
that of more exposed shores. Chu et al. (2000) found 
the opposite on Hong Kong shores. They hypothesized 
that wave-exposed shores have more microhabitats to 
be exploited than do quiet bays. Our data do not sup- 


420 


port Chu’s observations, but our data suffer from un- 
even sampling in the various exposure categories and 
may not be as reliable. 

Light influences species composition especially on 
the more terrestrial shoreline habitats (Fletcher 1973a). 
For example, Caloplaca and Xanthoria species are 
almost entirely restricted to sunny rocks in Gwaii 
Haanas. Fletcher (1973a) found that most lichens in 
the Littoral Fringe and even the Lower Supralittoral 
Zone are largely indifferent to light conditions, but 
Ryan (1988b) reported that Verrucaria mucosa only 
occurs in shaded habitats. This species was found in 
both unshaded and shaded sites in Gwaii Haanas, 
although it was more abundant on shaded limestone. 
Since exposure to sun affects moisture availability and 
retention, some correlation of species distribution ac- 
cording to shade conditions is to be expected. Shaded 
shores are almost always also protected shores, so it 
is difficult to separate the two factors in explaining 
differences in lichen vegetation. 

Shoreline rocks are frequently subject to manuring 
by sea birds, increasing the nitrogen load (urea and 
ammonia) and raising the pH of the rock surface, and 
this affects the composition of the saxicolous lichen 
communities, especially in the salt spray zone. Orni- 
thocoprophilous lichens that prefer such habitats in- 
clude Lecanora straminea, Caloplaca verruculifera, 
Physcia caesia and Xanthoria candelaria, all present 
on the shores of Gwaii Haanas wherever the conditions 
are suitable. 

The relative position of the marine and maritime 
species in Gwaii Haanas agree closely with observa- 
tions made by others, in so far as there are species in 
common, e.g., Verrucaria mucosa, V. striatula, Pyreno- 
collema halodytes occur in the upper edges of the in- 
tertidal zone; V. maura appears higher and has a broad 
distribution; Xanthoria candelaria, Physcia caesia and 
Caloplaca verruculifera are associated with bird rocks; 
etc. (Eliasson 1965; Ryan 1988b; Sheard and Ferry 
1967; Sheard 1968; Fletcher 1973a; Fletcher 1973b). 

The west coast endemics, and especially the mari- 
time lichens closely restricted to Haida Gwaii, com- 
prise 17 of the 67 species listed in the Appendix. It is 
useful to examine these species to see what their eco- 
logical and taxonomic equivalents would be elsewhere 
in the world. We will begin with ecological relation- 
ships. 

Caloplaca litoricola, a Haida Gwaii specialty, is in 
the Supralittoral Caloplaca zone as one would expect, 
but, since it is grey rather than yellow, it does not help 
to create an orange zone as other species of Caloplaca 
do in Europe or even eastern North America. On the 
other hand, the Pacific Northwest endemics C. rosei 
and C. inconspecta, and especially Caloplaca flavo- 
granulosa, are the ecological equivalents of the similar 
European C. marina (also in western North America 
from California to Vancouver Island, but not reaching 
Haida Gwaii; Arup 1992), which is part of the “orange 
zone” of European shores. Instead of the fruticose 


THE CANADIAN FIELD-NATURALIST 


Vol. 117 


cyanophilic lichen, Lichina pygmaea, common in the 
British Isles (and rare on the American east coast), 
we have Spilonema revertens, a microfruticose cyano- 
philic lichen in Gwaii Haanas, although it tends to be 
higher in the transects than Lichina would appear. In- 
stead of species like the orange foliose lichen, Xan- 
thoria parietina (common in both Europe and along 
the east coast of North America) in the Middle to 
Upper Supralittoral Zone, Haida Gwaii has the white 
crustose lichen Coccotrema maritimum, resulting in 
a conspicuous white zone replacing the orange zone 
seen in Europe. We have no ecological equivalent for 
their maritime saxicolous fruticose lichen, Ramalina 
siliquosa, but instead of Fuscidea cyathoides, Haida 
Gwaii has the narrowly endemic F: thomsonii in the 
Terrestrial zone (Brodo and Wirth 1998). 

The distribution of geographically or ecologically 
separated, closely related taxa (vicariants) with respect 
to these endemics or near-endemics is especially in- 
teresting. The categories of relationships and the taxa 
involved are summarized below, in most cases, with 
references to discussions already published. 


High Mountain connections 

The following species either have close relatives in 
the arctic-alpine flora, or have populations/subspecies 
that are found only on maritime and alpine rocks, not 
in between. 

Caloplaca litoricola: C. exsecuta (Ny1.) Dalla Torre 
& Sarnth. (Brodo 1984) 

Ochrolechia subplicans: subsp. hultenii 1s on shore- 
line rocks; subsp. subplicans grows in the alpine zone 
from Haida Gwaii to Alaska (Brodo 1988). 

Porpidia carlottiana: populations on Haida Gwaii 
either on shore or in mountains; close to P. glauco- 
phaea (Korber) Hertel & Knoph in Hertel, an ocean- 
ic species (Gowan 1989). 


Southern Hemisphere connections 

These species are closely related to species common 
in the southern hemisphere, either in South America 
or Australasia. 

Coccotrema maritimum: C. cucurbitula common on 
trees in Chile, etc. as well as in other parts of the Paci- 
fic Basin, e.g., Japan (Brodo 1973) 

Collema fecundum: C. novozelandicum Degel., on 
calcareous non-maritime and maritime rocks and soil, 
South Island, New Zealand (Degelius 1974, 1979) 

Verrucaria epimaura: V. durietzii 1. M. Lamb, mari- 
time lichen from the Aukland Islands (Brodo and 
Santesson 1997) 

Kohlmeyera complicatula: the same or related spe- 
cies are On maritime rocks in western South America. 
(Brodo 1976) 


Related to widespread maritime species 

These species are most closely related to other mari- 
time lichens. 

Lecanora species number 1: L. actophila, L. heli- 
copsis This undescribed Lecanora with a thick, white, 
xanthone-containing thallus will be discussed in a pub- 


2003 


lication in preparation. It is clearly related to other 
species in the L. dispersa aggregate such as those 
listed above. 

L. poliophaea (undescribed subspecies): subsp. 
poliophaea. The Haida Gwaii populations of L. 
poliophaea differ in some respects from European 
populations and are probably deserve taxonomic 
recognition. This will be discussed in the publication 
in preparation mentioned above. 

Verrucaria. schofieldii: V. erichsenii, a widespread 
supralittoral species (Brodo and Santesson 1997). 


Related to widespread non-maritime species 

The following species have their closest relatives 
as Temperate or Boreal, non-maritime lichens. 

Fuscidea thomsonii: F. intercincta (Nyl.) Poelt, F: 
atlantica (H. Magn) P. James & Poelt, both are Euro- 
pean species found in regions with an oceanic climate 
(Brodo and Wirth 1998; Oberhollenzer and Wirth 1984). 

Fuscopannaria maritima: F. thiersii P. M. J@rg., a 
Pacific Northwest species on iron-rich moist rocks 
(Jérgensen 2000); and F: leucostictoides Ohlsson, a 
western American corticolous species (Jorgensen 1978). 

Porina pacifica: Porina chlorotica (Ach.) Miill. Arg., 
widespread northern species on shaded rock faces 
(Brodo 2004). 

Rhizocarpon maritimum ined: R. cinereovirens is a 
Temperate species on shaded and exposed rocks, B. C., 
Black Hills, Long Island, N.Y. (The species, which will 
be described in a forthcoming paper, is closely related 
to R. cinereovirens (Fryday 2002: 463). 

These phytogeographic relationships of the endemic 
or near-endemic maritime lichens of Haida Gwaii and 
their vicariant taxa suggest survival in the area during 
the Pleistocene glaciations. Relationships with montane 
species may indicate, as has been suggested (Clague 
1989; Heusser 1989), that during the several glacial 
advances of the Pleistocene, some coastal areas and 
the highest mountain peaks of Haida Gwaii remained 
exposed. They would, therefore, have been available 
for colonization by lichens. It is possible that species 
that evolved at sea level from montane ancestors were 
highly restricted to coastal rocks and never were able to 
reinvade the intervening elevations although the highly 
exposed and harsh seacoast environment has much in 
common with alpine habitats and often shares the same 
or related species. It is probably significant that the 
closest relative of Coccotrema maritimum, a rock- 
dwelling species, is C. cucubitula, primarily a corti- 
colous species throughout its range. A glacial relic 
would, of course, have to survive on non-arboreal sub- 
strates. Schofield (1989) notes that all the Haida Gwaii 
endemics among the bryoflora occur on rock or soil. 

Many organisms are distributed around the Pacific 
Rim, or are disjunct from regions thousands of kilo- 
metres away. Plants and animals showing these pat- 
terns usually are related at the genus level or above, 
but these relationships show up at the species level in 
lichens and sometimes bryophytes (Schofield 1989). 


BRODO AND SLOAN: LICHEN ZONATION ON COASTAL ROCKS 


The fact that the land mass of Haida Gwaii represents 
a crustal plate fragment (terrane) that originated off the 
coast of Peru and and started its northward migration 
approximately 230 million years ago (Sutherland- 
Brown and Yorath 1989) provides a possible way for 
the species to have been transported (assuming the spe- 
cies had evolved by that time), although long distance 
dispersal is another means. Similar patterns are known 
from bryophytes; e.g., the moss Polytrichadelphus 
lyellii Mitt, which is known from British Columbia 
to California. The other 20 species of the genus are 
from South America, New Zealand and Australia 
(Persson, H. 1968. The possible migration routes of 
the species with southern hemisphere affinities are 
discussed by Brodo (1973). 

Maritime lichens on Haida Gwaii that are disjunct 
with western Europe or even eastern North America 
probably reveal ancient, probably Tertiary, distribu- 
tion patterns that were at first continuous but became 
dissected by climatic and geological events over the 
past 10-20 million years leaving the Haida Gwaii 
populations clinging to their outposts in microclimat- 
ically suitable habitats. Equivalent patterns among 
the mosses are common (Schofield 1984). 


Conclusions 

Rock-dwelling marine lichens of Gwaii Haanas are 
distributed in zones much as they are elsewhere in the 
world, although some species found abundantly only in 
Haida Gwaii create a unique appearance of these zones, 
such as the white Coccotrema zone, as compared with 
other regions. The unusually large percentage of en- 
demic, near-endemic or disjunct species suggests that 
at least the headland rocks along the west coast were 
refugia during the last glacial maximum. 


Acknowledgments 

We thank Gwaii Haanas (Parks Canada) for fund- 
ing the summer 2000 fieldwork and Doug Burles and 
Lee Edenshaw for field assistance. Dr. Fenja Brodo 
made numerous helpful suggestions on early drafts and 
assisted with some quadrat studies when not execut- 
ing her own Gwaii Haanas inventory project. IMB, 
now retired from the Canadian Museum of Nature, 
thanks the Museum for appointing him as a Research 
Associate and providing him with space and facilities 
for his lichen research. We dedicate this paper to the 
memory of Bruce Ryan, who first studied the maritime 
zonation of lichens in the Pacific Northwest and did 
it so well. 


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Received 30 June 2003 
Accepted 17 February 2004 


424 


THE CANADIAN FIELD-NATURALIST 


Volany 


Appendix: Lichens of Haida Gwaii Found on Maritime Rocks 


Note: Many of the lichens listed below are non-maritime lichens that, on occasion, are found growing in the maritime 
zone thereby demonstrating a tolerance for salt spray. Species not found in Gwaii Haanas National Park Reserve are marked 
with an asterisk (*). Those new to British Columbia are in boldface. Voucher specimens of all these records can be found in 


the Canadian Museum of Nature (CANL). 


Adelolecia koalensis (Ny1.) Hertel & Rambold (Syn. Lecidea 
conferenda Nyl.) This was first reported for British Col- 
umbia in Brodo (1995). 

*Amandinea coniops (Wahlenb. in Ach.) M. Choisy ex Scheid. 
& H. Mayrh. 

Arthonia phaeobaea (Norman) Norman 

Aspicilia caesiocinerea (Nyl. ex Malbr.) Arn. 

*A spicilia supertegens Arn. This is not normally a maritime 
species. The single voucher specimen requires further study. 

*Bacidina inundata (Fr.) Vézda. This species was also found 
in streams in Haida Gwaii. 

*Caloplaca chlorina (Flotow) H. Olivier. The voucher speci- 
men was verified by both T. Tonsberg and C. M. Wetmore. 

Caloplaca citrina (Hoffm.) Th. Fr. 

Caloplaca flavogranulosa Arup 

*Caloplaca inconspecta Arup 

Caloplaca litoricola Brodo 

Caloplaca rosei Hasse 

Caloplaca verruculifera (Vainio) Zahlbr. 

*Catillaria chalybeia (Borrer) Massal. 

*Cliostomum griffithii (Sm.) Coppins 

Coccotrema maritimum Brodo 

Collema fecundum Degel. 

*Ephebe lanata (L.) Vainio 

*Fuscidea mollis (Wahlenb.) V. Wirth & Vézda. The voucher 
specimen is not typical of the species, so this record is 
questionable. 

Fuscidea thomsonii Brodo & V. Wirth 

Fuscopannaria maritima (P. M. Jorg.) P. M. Jorg. 

Gyalecta jenensis (Batsch) Zahlbr. 

Herteliana alaskensis (Nyl.) Zahlbr. 

*Kohlmeyera complicatula (Nyl.) Schatz 

Lecanora muralis (Schreber) Rabenh. 

*Lecanora poliophaea (Wahlenb.in Ach.) Ach. s. lat. 

Lecanora straminea Wahlenb. ex Ach. 

*Lecanora tenera (Nyl.) Cromb. 

Lecanora sp. number 1. This is a xanthone-containing. mari- 
time, saxicolous member of the Lecanora dispersa group 
and almost certainly is undescribed. 

Lecanora sp. number 5. This lichen is similar in many respects 
with L. contractula Ny\l. 

*Lecidella scabra (Taylor) Hertel & Leuckert 

Lecidella stigmatea (Ach.) Hertel & Leuckert f. stigmatea 

Ochrolechia subplicans (Ny1.) Brodo subsp. hultenii (Erichs.) 
Brodo 

Opegrapha gyrocarpa Flotow 

Opegrapha sp. This is a sorediate species containing roccel- 
lic acid, with 3-septate spores and lirelliform ascomata 
unlike those of O. gyrocarpa. It may be undescribed. 

Parmelia saxatilis (L.) Ach. 


Physcia caesia (Hoffm.) Fiirnr. 

Placopsis lambii Hertel & Wirth. There is a single record of 
P. gelida (L.) Lindsay s. str. on a rock in a tidal marsh, but 
otherwise the species is only known on Haida Gwaii from 
inland localities, usually on roadsides and fresh rock 
exposures. 

*Polychidium muscicola (Sw.) S. F. Gray 

Porina pacifica Brodo. This species, which superficially 
resembles P. chlorotica (Ach.) Miill. Arg., was only recent- 
ly described (Brodo 2004) 

Porpidia carlottiana Gowan 

Porpidia contraponenda (Arnold) Knoph & Hertel 

Porpidia speirea (Ach.) Kremp. 

Porpidia thomsonii Gowan 

*Punctelia stictica (Duby) Krog 

Pyrenocollema halodytes (Nyl.) R. C. Harris s. lat. We have in- 
cluded specimens with sunken perithecia as well as those 
with sessile perithecia under this name if the perithecia 
were under 0.3 mm in diameter. The much larger P. sub- 
litorale (Leighton) R. C. Harris ex Fletcher in Coppins, 
P. James & D. Hawksw. has not yet been found in Haida 
Gwail. 

Rhizocarpon geminatum Korber 

Rhizocarpon “haidensis” Brodo, ined. This species is closely 
related to R. cinereovirens (Miill. Arg.) Vainio (Fryday 
2002). Its description is being prepared for publication. 

*Rhizocarpon hensseniae Brodo 

Rhizocarpon hochstetteri (K6rber) Vainio s. str. A detailed 
discussion of this species is provided by Fryday (2002). 

*Rinodina gennarii Bag}. 

Spilonema revertens Ny. 

*Tephromela atra (Hudson) Hafellner in Kalb 

*Tylothallia biformigera P. James & R. Kilias in R. Kilias 

Verrucaria amphibia Clemente 

Verrucaria degelii R. Sant. 

Verrucaria epimaura Brodo 

Verrucaria erichsenii Zsch. 

*Verrucaria halizoa Leighton 

Verrucaria maura Wahlenb. in Ach. 

Verrucaria mucosa Wahlenb. in Ach. 

Verrucaria sandstedei de Lesd. This species, like V. striatula 
listed below, was reported from coastal rocks in Fildago 
Island, Washington, by Ryan (1988a, b). 

Verrucaria schofieldii Brodo 

*Verrucaria Silicicola Fink 

Verrucaria striatula Wahlenb. See comment under V. sand- 
stedel. 

Verrucaria sp. number 5 (dry rock at edge of beach) 

Xanthoria candelaria (L.) Th. Fr. 


Predaceous Water Beetles (Coleoptera: Adephaga: Dytiscidae, Gyrinidae) 
Collected Along the Horton and Thelon Rivers in the Arctic Central 
Barrens of Canada 


HELENA V. SHAVERDO! and DONNA J. GIBERSON? 


‘Department of Entomology, 213 Animal Science/Entomology Building, University of Manitoba, Winnipeg, Manitoba R3T 2N2 
Canada 
"Department of Biology, University of Prince Edward Island, Charlottetown, Prince Edward Island C1A 4P3 Canada 


Shaverdo, Helena V., and Donna J. Giberson. Predaceous water beetles (Coleoptera: Adephaga: Dytiscidae, Gyrinidae) col- 
lected along the Horton and Thelon rivers in the Arctic Central Barrens of Canada. Canadian Field-Naturalist 
118(3): 425-433. 


Predaceous water beetles were collected during expeditions along two northern Canadian rivers during 2000 and 2002. 
Twelve species of Dytiscidae (including 11 named species and one additional genus identified from a larva) and one species of 
Gyrinidae are recorded from 20 sites along the Horton and Thelon rivers in the Central Barrens area of the Canadian Arctic. 
These records represent an extension of the distributions of four species to the northeast in the Northwest Territories (NWT), 
and two species to the northwest in Nunavut (NU). Oreodytes sanmarkii is reported for NWT and for the mainland of NU 
for the first time. //ybius erichsoni, Hydroporus geniculatus, and Gyrinus opacus are reported for NU for the first time. Five 
species were recorded for the first time from the Southern Arctic ecozone, and one from the Taiga Shield ecozone. The 
majority of specimens were collected in habitats that were consistent with those previously known for each species. 


Key Words: Insecta, Coleoptera, Adephaga, Dytiscidae, Gyrinidae, predaceous water beetles, faunistics, river, Northwest 


Territories, Nunavut, Arctic Central Barrens. 


The Central Barrens area of the Canadian Arctic is 
a vast and very sparsely populated area dominated by 
arctic tundra, scattered clumps of taiga vegetation, 
large lakes, and rivers. The inaccessibility of the region 
has meant that the area has been poorly collected for 
all insect groups (Currie et al. 2000), including preda- 
ceous water beetles. Only three surveys have been con- 
ducted on aquatic Adephaga in the Yukon, Alaska, 
and the eastern Canadian Arctic (Alarie and Maire 
1991; Larson 1991, 1997), though data on this group 
of Coleoptera in arctic and subarctic regions can also 
be found in Larson and Roughley (1991), Roughley 
(1991), Anderson (1997), and Larson et al. (2000). 
Recently a series of expeditions were mounted to ex- 
plore the aquatic insect fauna along large river systems 
on the treeless barrens of the mainland Canadian Arc- 
tic zone (Currie et al. 2000, 2002). The objective of 
this paper is to report on the predaceous water beetles 
collected from habitats in and near the Horton River 
Northwest Territories (NWT) and the Thelon River, 
NWT and Nunavut (NU), and present faunistic and 
habitat data from the collections. 


Methods 

Two expeditions were made by DJG and colleagues 
from various institutions to northern Canadian rivers 
during 2000 and 2002 (Figure 1). Expedition mem- 
bers flew in to upstream sections of the Horton and 
Thelon rivers in NWT and travelled downriver by 
canoe, stopping frequently to sample habitats along the 
linear corridor of each river. Aquatic sampling was 


carried out using a D-frame kick net with a 200 um 
mesh. In flowing water habitats, the net was placed 
into the stream and the substrate disturbed by kicking 
or by moving rocks by hand, and the dislodged insects 
were carried into the net by the current. In areas with 
little or no flow, the substrate was disturbed as above, 
and then the net was drawn through the water by 
hand to catch any dislodged insects. Insects were 
sorted alive from the substrate materials on site, and 
preserved in 80% ethanol. Predaceous water beetles 
were not specifically targeted in the study, but those 
that were collected (61 adults and 6 larvae) were exam- 
ined by HVS. Collection records were compared to the 
maps and distribution table in Larson et al. (2000) to 
determine whether any of them represented range ex- 
tensions among the provinces and ecozones. The doc- 
ument A National Ecological Framework for Canada 
(Ecological Stratification Working Group 1995) was 
consulted for the classification of Canada’s ecological 
regions. Collecting Permits were obtained from the 
Aurora Research Institute (Inuvik, NWT) and the 
Nunavut Research Institute (Iqaluit, NU) to carry out 
the research. Voucher specimens have been deposited 
in the collections of the Wallace Museum (University 
of Manitoba) and the Royal Ontario Museum, and in 
the private collection of D. J. G. 


Study Site and Sampling Regime 

The Horton River begins west of Horton Lake, which 
is located north of Great Bear Lake in the Northwest 
Terriories of Canada (Figure 1). The river, which has 


425 


426 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Horton River 


68°. 


127° Tanptalase 123° 121° 


Thelon River 


Thelon Game" 
* Sanctuary \ 


ynaeunn 


_souloyuse| ISOMYYON 


105°W 100°W 


FiGureE 1. Location of study area (A), including the Horton River (B) and Thelon River (C) showing sample sites. The fine 
dashed line indicates the watershed boundary upstream of the lowest sampled site for both watersheds. The grey 
border on the Thelon Watershed map indicates the boundaries of the Thelon Game Sanctuary. Site co-ordinates are 


given in Table 1. 


a drainage basin area of approximately 26 680 km? 
and a total length of = 800 km, runs entirely in the 
Southern Arctic Ecozone. Expedition members col- 
lected over a 700 km stretch of the river from Horton 
Lake to the Arctic Ocean. The river flows over lime- 
stone-dominated substrates in its upper reaches, where 
it is characterized by relatively high pH (>8.0) and spe- 
cific conductance (>150 uS/cm). In its lower reaches, 
the river flows through the Smoking Hills, a region 
containing lignite and magnesium deposits which spon- 
taneously ignite when exposed to air (for example, 
during bank slumping during summer). These result 
in “smoke holes” as smoke pours from places where 
the coal is burning underground, and the water drain- 
ing these sites is profoundly affected by the geology. 
Tributaries and ponds in this region were highly acidic 
(some with pH values < 2.5), and acid runoff in this 
area resulted in a decline in river pH to approximately 
neutral values. The river is clear and swift-flowing for 
most of its length, but becomes muddy and meander- 
ing in its lower portion. Sampling was carried out in 
37 sites on or near the river between 17 July and 8 Au- 
gust 2000. Predaceous water beetles were collected 
from 5 of the 37 sites along the river, corresponding to 


sampling sites 8, 11, 14, 18, and 23 of the larger Hor- 
ton River survey; all were located in the middle sec- 
tion, which was clear and swift-moving (Figures 1b, 
2; see Table 1 for descriptions of each site). 

The Thelon River begins near Lynx Lake, to the 
east of Great Slave Lake (Figure 1). The river flows 
= 900 km from Lynx Lake to Baker Lake (which exits 
via Chesterfield Inlet to northern Hudson Bay to the 
east) and drains a total area of = 240 000 km?. The 
upper portions of the Thelon River are in the Taiga 
Shield ecozone, then the river flows into the Southern 
Arctic ecozone. Expedition members flew into the 
junction of the Hanbury and Thelon rivers, and col- 
lected along a 300 km stretch of the Thelon River 
located within the bounds of the Thelon Game Sanc- 
tuary (Figure Ic). The part of the river covered in this 
survey (i.e., upstream of Beverly Lake, NU) drains 
an area of 71 470 km. The river corridor upstream of 
the study area consists of large numbers of lakes, water- 
falls, and rapids, but in the study area itself, the river 
meanders over an ancient glacial lake bottom. At this 
point, the river is wide and slow moving and dominat- 
ed by sandy or sand/silt-embedded cobble substrates. 
Surrounding relief is very low except in a few spots, 


2004 


Horton Site 8 
one 


SHAVERDO and GIBERSON: PREDACEOUS WATER BEETLES IN ARCTIC 


Horton Site 11 


FIGURE 2. Study sites along the Horton River where predaceous water beetles were collected, July/August 2000 (compare to 
Figure 1B for site locations). 


where low hills, boulders, and more rapid flow can be 
found. Most of the river flows through open tundra, 
but a small stretch in the upper third of the study area 
(= 65 km) is located in a remnant spruce and larch 
forest known as the “Thelon Oasis”. The pH and spe- 
cific conductance values were quite low, especially in 
comparison with the Horton River, with pH values gen- 
erally around 6.0, and specific conductance values 
= 10 uS/cm for main river sites. pH and conductivity 
values for tributaries and tundra pools were more 
variable, however. Sampling was carried out at 34 sites 
on or near the river between between 29 June and 11 
July 2002. Predaceous water beetles were collected 


from 15 of the 34 sites, representing a combination 
of river and pool sites corresponding to sites 1, 3, 4, 
Gy TS OMI lor slSeil9) 22-26::27.) 50and 32-oftthe 
larger Thelon River survey (Figures Ic, 3; see Table 
1 for descriptions of each site). 


Results and Discussion 

Oreodytes laevis (Kirby) and O. sanmarkii (Sahlberg) 
were the two most common species found in our north- 
em rivers survey, and were also the only species found 
in both rivers (Table 1). Oreodytes laevis accounted 
for 26 of the 61 adults collected (> 40%) and was found 
in eight of the 20 sites where water beetles were found. 


428 THE CANADIAN FIELD-NATURALIST Vol. 118 


_ Thelon Site 7 


FiGuRE 3a. Study sites along the upper reaches of the Thelon River where predaceous water beetles were collected, July/ 
August 2000 (compare to Figure 1C for site locations). 


2004 SHAVERDO and GIBERSON: PREDACEOUS WATER BEETLES IN ARCTIC 429 


Thelon Site 18 i Thelon Site 19 


yA 


Thelon Site 30 


FIGURE 3b. Study sites along the lower reaches of the Thelon River where predaceous water beetles were collected, July/August 
2000 (compare to Figure Ic for site locations). 


430 


TABLE |. Distribution and abundance of predaceous water beetles collected along the Horton and Thelon Rivers. All species 
were collected in the main branch of the rivers except where specifically indicated. All species were in the family Dytisci- 


THE CANADIAN FIELD-NATURALIST 


dae except Gyrinis opacus which was in the family Gyrinidae. 


Site 


Coordinates / Date Site Details 


Horton River, NWT 


8 


67°59°24.5"N 
123°13°55.5°W 
20 July 2000 


68°09°29"N 
123°22°44.9"W 
22 July 2000 


68°23°31.5°N 
123°38°03.4°W 
23 July 2000 


68 39°3.18°N 
124°08°53.0"W 
25 July 2000 
68°44’ 19.9"N 
124°59°0.8°W 
28 July 2000 


Thelon River, NWT 


1 


13 


16 


18 


63 38°04.8"N 
104°32°18.6"W 
29 June 2002 


63 42°04.4”"N 
104°25’26.6" W 
30 June 2002 


63 48’22°N 
104°18’21.1"W 
30 June 2002 


63°49’58.4"N 
104°06°20.9"W 
1 July 2002 


63°55°02.7"N 
103°56’59.5°W 
1 July 2002 


64°02’ 13.2”N 
103°51°41.8°W 
2 July 2002 


64°17°01.8"°N 
103°33°22.4”"W 
3 July 2002 


64°19" 18.6”N 
103°19715.7°W 
4 July 2002 


64°17°56.2”N 
102°43°20.6"W 
5 July 2002 


cobble dominated riffle and shore section, 
where river slowed and flowed through a 
broad valley; pH: 8.3; conductivity: 

150 wS/cm; temp.: 15°C 


slow, weedy shoreline area along 

the main channel of the river (fill 

in from logbook); pH: 8.2; 
conductivity: 180 uS/cm; temp.: 10.5°C 


backwater zone and a riffle, from a portion 
of the river in a narrow valley surrounded 
by high limestone cliffs; pH: 8.2; 
conductivity: 180 uS/cm; temp.: 12°C 


the river is a series of very shallow 
flats and riffles; temp.: 12°C 


canyon/white-water area of the river, deep 
bedrock pools and fast riffles; at camp spot 
no obvious riffles though water moving very 
fast; sampled from shore; temp.: 17°C 


relatively slow and wave washed section near 
shore, in silt/sand embedded cobble; pH: 6.2; 
conductivity: 10 uS/cm; temp.: 11°C 


rocky shore with fairly strong current, 
and in backwater pools behind large 
boulders; pH: 6.2; conductivity: 

10 wS/cm; temp.: 11°C 


slow section near shore, with cobble substrate 
embedded in sand and silt; pH: 6.2; 
conductivity: 10 uS/cm; temp.: 10.5°C 


submerged grasses in slow current near 
shore, 10 m upstream of a tributary entering 
from the south; pH: 6.2; conductivity: 

40 uS/cm; temp.: 14°C 


cobble and silt substrate near shore where 
main river narrowed and became swift 
flowing, even near shore; pH: 6.2; 
conductivity: 10 uS/cm; temp.: 11°C 


at Hornby Point; well embedded cobble 
in fast, shallow riffle near shore; pH: 6.1; 
conductivity: 10 uS/cm; temp.: 12°C 


cobble substrate in fast water in main 
channel, plus slower water near shore; 
pH: 6.3; conductivity: 30 uS/cm; temp.: 13°C 


unnamed tributary entering Thelon River from 


forest to the north; riffle with large clean cobble 


substrate; pH: 7.6; conductivity: 40 uS/cm; 
temp.: 16°C 


river wide and lake-like; wave-washed area, 
in cobble substrate well embedded in sand; 


pH: 6.3; conductivity: 10 uS/cm; temp.: 13.5°C 


Species 


Oreodytes laevis 
(Kirby) 

Oreodytes sanmarkii 
Sahlberg 


Oreodytes sanmarkii 


Oreodytes laevis 


Oreodytes sanmarkii 


Oreodytes sanmarkii 


Oreodytes sanmarkii 


Oreodytes laevis 


Oreodytes sanmarkii 


Oreodytes laevis 


Oreodytes laevis 
Hygrotus novemlineatus 
(Stephens) 


Oreodytes laevis 


Stictotarsus griseostriatus 
(DeGeer) 


Oreodytes laevis 


Stictotarsus griseostriatus 


Oreodytes sanmarkii 


Oreodytes laevis 


Vol. 118 


Specimens | 


1° 


ie) 


lo 


2c¢c 


lo 


ilxe} 


2c¢c 


oil © 


2004 


SHAVERDO and GIBERSON: PREDACEOUS WATER BEETLES IN ARCTIC 


43] 


TABLE | (continued). Distribution and abundance of predaceous water beetles collected along the Horton and Thelon Rivers. 
All species were collected in the main branch of the rivers except where specifically indicated. All species were in the fami- 
ly Dytiscidae except Gyrinis opacus which was in the family Gyrinidae. 


Site 


19 64°10°46.1°N 
102°36°58.1"W 
6 July 2002 


23 64°11°05.4"N 
102°19°06.9° W 
7 July 2002 


Thelon River, NU 

26 64°19713.2”°N 
101°50’11.1°W 
8 July 2002 


27 64°19’ 13.2”N 
101°50°11.1"W 
8 July 2002 


30 64°26°15°N 
101°43°55.1"W 
9 July 2002 


64°32°25.2”N 
101°24°48.9°W 
10 July 2002 


Coordinates / Date Site Details 


small brown-water tundra pools above the 

river on north, pools draining to each other 

and seeping to the river; overhanging vegetation, 
undercut banks, stony bottom; pH: 6.0; 
conductivity: 20 uS/cm; temp.: 14°C 


unnamed tributary seep entering the river 
from the north and small brown-water tundra 
pools surrounded by willows, shorelines with 
sphagnum and grass; pH: 5.7; conductivity: 
50 wS/cm; temp.: 9°C 


unnamed tributary entering the river from the 
north; cobble substrate in riffle area of the stream; 
pH: 6.5; conductivity: 10 uS/cm; temp.: 11°C 


small brown-water tundra pond beside site 26; 
much submerged and decaying vegetation (grass); 
pH: 6.0; conductivity: 10 uS/cm; temp.: 15°C 


large tundra pond on southern bank above the 
river, with overhanging willows and soft bottom 
substrates; pH: 6.0; conductivity: 10 uS/cm; 
temp.: 12°C 


unnamed small tributary entering the river 
from the south - small pools and seeps between 
the pools; overhanging vegetation; pH: 6.2; 
conductivity: 20 wS/cm; temp.: 14°C 


Species Specimens 
Hydroporus appalachius 2d¢0 
Sherman DIRE) 
Agabus confines 499 
(Gyllenhal) il bake 
Ilybius erichsoni 1E9 
(Gemminger 
and Harold) 
Agabus sp. DES 
Oreodytes sanmarkii 1° 
Hydroporus geniculatus lo 
Thomson 
Ilybius erichsoni lo 
2IE3e 
Stictotarsus griseostriatus lo 
Agabus arcticus ls 
(Paykull) 6 29 
Rhantus sp. WLR 
Gyrinus opacus Sahlberg le 
Hydroporus morio 3.99 
Aubé 
Agabus thomsoni IN Gr Ale. 
(Sahlberg) 


Note: * L3 — larva of third instar; L2 — larva of second instar. 


Eleven specimens of Oreodytes sanmarkii were also 
found at eight sites. Stictotarsus griseostriatus (De- 
Geer) specimens were collected in three of the sites, but 
there was only a single specimen in each collection. The 
remaining species were all restricted to a single site 
each. Although several specimens of individual species 
were often found, for example, the sites with Agabus 
confinis (Gyllenhal) and A. arcticus (Paykull) (Table 1). 

These collections are range extensions among the 
provinces and ecozones of Canada for most of the 
species collected, and represent a “filling in of gaps” in 
the distributions of many northern species. Because of 
logistic difficulties in sampling these interior regions 
of the arctic barrens, many previously reported species 
distributions have appeared disjunctive, or restricted 
to coastal areas (e.g., Larson et al. 2000). For example, 
the known ranges of Hydroporus appalachius Sher- 
man, Oreodytes laevis, and Agabus confinis are extend- 
ed to the northeast within the Northwest Territories, 
and those of Agabus arcticus and A. thomsoni Salh- 
berg are extended to the northwest within Nunavut. 


Oreodytes laevis and Hydroporus appalachius are 
widespread in the boreal low arctic zone of the Nearc- 
tic, so these records represent an extension into the 
Tundra zone. With respect to ecozone distribution, O. 
laevis, H. appalachius, and A. confinis are recorded 
for the first time from the Southern Arctic ecozone, 
and A. confinis is now known from all ecozones of 
Canada except the prairies. [/ybius erichsonii (Gem- 
minger and Harold) is reported for the first time for 
Nunavut, and its known range is extended to the north- 
east within the Northwest Territories, and into the 
Southern Arctic Ecozone. Ilybius erichsonii has now 
been reported from all the ecozones of Canada. Hydro- 
porus geniculatus Thomson is a Holarctic species 
which occurs in the boreal and alpine zones in north- 
western North America. In Canada, it is distributed 
throughout most of British Columbia and western 
Alberta, and north into the Yukon Territory and Alaska 
(Larson et al. 2000), and is reported for the first time 
in this study in Nunavut and in the Southern Arctic 
ecozone. The present record is an extension of its known 


432 


distribution far to the northeast, and therefore the 
range for this species appears disjunctive. However, 
H. geniculatus belongs to the poorly studied nigellus- 
group, and revision of the group is needed to clarify 
its taxonomic position and distribution. We report the 
presence of Oreodytes sanmarkii in the Northwest 
Territories and in the Taiga Shield ecozone for the first 
time, and the collections along the Thelon River also 
represent the first record of the species on the Nunavut 
mainland. This Holarctic species was previously known 
in Canada from a few sites in northern Yukon and a site 
on Southampton Island (NU) (Larson et al. 2000). 
Hygrotus novemlineatus (Stephens), Hydroporus morio 
(Aubé), and Stictotarsus griseostriatus have been 
reported in the Canadian Barrens to the east and west 
of the sampled areas, but these collections indicate 
that their distributions are more continuous across the 
north than previous distribution maps might indicate. 
Gyrinus opacus (Gyrinidae) is reported for Nunavut 
for the first time, and collections on the Thelon River 
may represent the most northeasterly record for the 
species (Oygur and Wolfe 1991). However, this record 
is in need of confirmation since it is based on study 
of a single female specimen. 

Most of the species were collected in habitats that 
matched those that were previously reported for each 
species (e.g., Larson et al. 2000). However, this study 
has produced some new and additional data on habi- 
tat distributions for several species. Oreodytes laevis 
and O. sanmarkii have been reported in rivers, streams, 
and along exposed shorelines of cold lakes with little or 
no submerged vegetation, and with mineral substrates. 
These two species dominated in these types of habi- 
tat along the Thelon and Horton rivers. According to 
Alarie and Maire (1991; as H. hudsonicus) and Larson 
et al. (2000), Hygrotus novemlineatus is circumpolar in 
the low arctic region and has been collected in Canada 
(near Churchill Manitoba, and in Quebec) in brackish 
splash pools (rock-pools) along the coast. However, 
in our survey it was collected among dense submerged 
grass in slow water current along the shore of the 
Thelon River. The species may have relatively wide 
habitat preferences, since Nilsson and Holmen (1995) 
reported it in Fennoscandia from similar habitats to 
the Thelon site, as well as from sandy oligotrophic lake 
margins and sparsely vegetated ponds. In the southern 
parts of its range, Hydroporus appalachius is usually 
found in habitats where there is some water movement, 
for example in streams, springs, and the margins of 
small lakes (Larson et al. 2000). In Labrador, the 
species was collected from a protected shoreline of a 
large lake (Larson et al. 2000), and it has been found 
in inland pools in the Quebec subarctic region (Alar- 
ie and Maire 1991). The habitat in the more northern 
parts of its range has not previously been described, 
and in this survey, it was found in small tundra pools 
that drained into each other before spilling into the 
Thelon River. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Acknowledgments 

We are grateful to Rob Roughley and Lisa Purcell 
for their advice and comments on the manuscript. 
Financial support for the collections was provided by 
an NSERC operating grant, a UPEI Senate Research 
Grant, and a Nature Discovery Grant (National Muse- 
ums of Canada) to D. J. Giberson. Lisa Purcell helped 
to collect the specimens. Support for the study of the 
material was provided by NSERC through a 2001 
NATO Science Fellowship to H. V. Shaverdo and an 
NSERC operating grant to Rob Roughley. Voucher 
specimens have been deposited in the J. B. Wallace 
Museum of the University of Manitoba, the Royal 
Ontario Museum, and in the private collection of 
H. V. S. Collecting permits were provided by the terri- 
torial governments through the Aurora Research Insti- 
tute in the Northwest Territories and the Nunavut 
Research Institute in Nunavut. 


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terists Bulletin 45: 280-284. 

Larson, D. J. 1997. Dytiscid Water Beetles (Coleoptera: 
Dytiscidae) of the Yukon. Pages 491-522 in Insects of the 
Yukon. Edited by H. V. Danks and J. A. Downes. Biological 
Survey of Canada. Monograph series (2), Biological Sur- 
vey of Canada (Terrestrial Arthropods). Ottawa. Ontario. 

Larson, D. J., and R. E. Roughley. 1991. Family Dytiscidae. 
Pages 62-72 in Checklist of beetles of Canada and Alaska. 
Edited by Y. Bousquet. Agriculture Canada Publication 
1861/E. 

Larson, D. J., Y. Alarie, and R. E. Roughley. 2000. Preda- 
ceous Diving Beetles (Coleoptera: Dytiscidae) of the Nearc- 
tic Region, with emphasis on the fauna of Canada and Alaska. 
NRC Research Press, Ottawa, Ontario, Canada. 982 pages. 

Nilsson, A. N., and M. Holmen. 1995. The aquatic Adephaga 
(Coleoptera) of Fennoscandia and Denmark. II. Dytiscidae. 
Fauna Entomologica Scandinavica 32: 1-192. 


2004 SHAVERDO and GIBERSON: PREDACEOUS WATER BEETLES IN ARCTIC 433 


Oygur, S., and G. W. Wolfe. 1991. Classification, distribution, Roughley, R. EK. 1991. Family Gyrinidae. Pages 72-73 in 
and phylogeny of North American (north of Mexico) spe- Checklist of beetles of Canada and Alaska. Edited by Y. 
cies of Gyrinus Miiller (Coleoptera: Gyrinidae). Bulletin Bousquet. Agriculture Canada Publication 1861/E. 
of the American Museum of Natural History 207: 1-97. 


Received 2 June 2003 
Accepted 14 July 2004 


Notes 


Collapsing Burrow Causes Death of a Eurasian Beaver, Castor fiber 


Liat R. THOMSEN!, FIONA SHARPE2, and FRANK ROSELL? 


' Department of Zoology, University of Aarhus, Denmark 


2 School of Biology, University of St Andrews, Fife, Scotland 


3 Faculty of Arts and Sciences, Department of Environmental and Health Studies, Telemark University College, N-3800 Bo 


i Telemark, Norway 


Thomsen, Liat R., Fiona Sharpe, and Frank Rosell. 2004. Collapsing burrow causes death of a Eurasian Beaver, Castor 


fiber. Canadian Field-Naturalist 118(3): 434-435. 


The death of a Eurasian Beaver Castor fiber caused by a collapsing burrow in southeast Norway is reported. Two days of 
heavy rainfall had presumably caused the burrow to collapse, suffocating the animal. 


Key Words: Beaver, Castor fiber, collapsing burrow, southeast Norway. 


Several causes of death have been reported for 
Eurasian Beavers (Castor fiber) (Rosell et al. 1996; 
Nolet et al. 1997), but to our knowledge this is the 
first report of a beaver killed due to a collapsing bur- 
row. The beaver was an adult solitary male (21.5 kg 
and 5 years old; age was determined by examining 
annual cementum and dentine layers of the first molar 
(van Nostrand and Stephenson 1964)). His mate died 
two months earlier, probably due to old age (18 years 
old). The animal was one of several radio tagged 
beavers that were followed during a field study in 
spring and summer 2000 in Telemark County, south- 
east Norway (59°25'N, 09°03'E) (see Campbell 2000). 
On the evening of 14 July 2000 it was noted that the 
pulse interval of the radio signal had increased. The 
signal is inversely related to the body temperature of 
the animal (Alterra 1999), thus a higher pulse interval 
indicated that the beaver had died. The dead beaver 
was located in the main part of a partly collapsed bur- 
row (assessed to be relatively new), which was exca- 
vated four days later in order to find and retrieve the 
carcass (Figure 1). 

Beavers dig burrows where banks are sufficiently 
high (Wilsson 1971; Zurowski 1992). A burrow usu- 
ally consists of a single living chamber, a water basin, 
and a tunnel with exit below the water level (Wilsson 
1971). The beaver was lying in the chamber (1 m long 
and 70 cm wide) of the collapsed burrow, facing to- 
wards the exit. The chamber was situated 210 cm from 
the water’s edge, 80 cm below the surface of the 
riverbank, and 60 cm above the current water level. 
Apparently the ceiling of the chamber had collapsed 
on top of the beaver causing the death of the animal, 
presumably by suffocation. There was loose sand along 
the flanks of and underneath the body of the dead 
beaver. The last 10 cm of the beaver’s tail was bent 
downwards into the sand and the right hind foot was 
crouched as if the beaver had been attempting to dig 
with it. The beaver had no external injuries and seemed 
to be in good condition. 


of a partly collapsed burrow. 


The burrow was dug in sandy soil. In the week prior 
to the discovery of the beaver there had been two days 
with heavy rainfall in the area with 30.8 mm and 24.6 
mm of rain, respectively (data from the Norwegian 
Meteorological Institute). 

We conclude that the combination of heavy rainfall 
and a sandy soil had caused the burrow to collapse, 
therefore causing the death of the beaver. How preva- 
lent this cause of death is in beavers is unknown. How- 
ever, we expect that more field studies using radio 
telemetry, in areas where beavers dig burrows, could 
clarify this issue. 


434 


2004 


Acknowledgments 

We thank Frode Bergan for help with excavating 
the beaver and Inger Hanssen-Bauer for providing the 
weather data. The study was financially supported by 
Telemark University College. The experiments comply 
with the current Norwegian law, the country in which 
they were performed. 


Literature Cited 

Alterra. 1999. Alterra (IBN/DLO) P.O. Box 23. 6700 AA 
Wageningen. Netherlands. 

Campbell, R. D. 2000. Territoriality in the European Beaver, 
Castor fiber. MSc thesis, School of Biological Sciences, 
University of East Anglia, Norwich, England. 

Nolet, B. A., S. Broekhuizen, G. M. Dorrestein, and K. 
M. Rienks. 1997. Infectious diseases as main factors of 


NOTES 


435 


mortality to beavers Castor fiber after translocation to the 
Netherlands. Journal of Zoology, London 241: 35-42. 

Rosell, F., H. Parker, and N. B. Kile. 1996. Causes of mor- 
tality in beaver (Castor fiber & canadensis). Fauna 49: 
34-46 [In Norwegian with English summary]. 

van Nostrand, F. C., and A. B. Stephenson. 1964. Age deter- 
mination for beavers by tooth development. Journal of 
Wildlife Management 28: 430-434. 

Wilsson, L. 1971. Observations and experiments on the ethol- 
ogy of the European beaver (Castor fiber L.). Viltrevy 8: 

_ E266: 

Zurowski, W. 1992. Building activity of beavers. Acta 
Theriologica 37: 403-411. 


Received 26 November 2002 
Accepted | November 2004 


Frequency of Tail Breakage of the Northern Watersnake, Nerodia 


sipedon sipedon 


KENNETH D. BOWEN! :2 


'Department of Biology, Central Michigan University, Mt. Pleasant, MI 48859 USA 


Present Address: 709 Ringold Street, Boone, Iowa 50036 USA 


Bowen, Kenneth D. 2004. Frequency of tail breakage in the Northern Watersnake, Nerodia sipedon sipedon. Canadian 
Field-Naturalist 118(3): 435-437. 


I noted the frequency of broken tails of Northern Watersnakes, Nerodia sipedon sipedon, in the Beaver Archipelago of North- 
eastern Lake Michigan. Overall, 10% (22 of 220) of captured snakes had broken tails. This value is similar to published values 
for closely related snakes but is smaller than those reported for another Nerodia sipedon sipedon population. Unlike some 
previously published studies, the frequency of injured tails was not greater for females or lesser for first-year snakes. The mech- 
anism behind the injury frequency observed here and the reason for differences between this and other studies are unknown. 


Key Words: Beaver Archipelago, Nerodia sipedon sipedon, Northern Watersnake, tail breakage, Michigan. 


The frequency of tail injuries in a lizard or snake 
population was at one time considered to be a useful 
index of the predation pressure on that population be- 
cause tail autotomy and breakage (Mendelson 1992) 
are thought to be important defense mechanisms (re- 
viewed in Arnold 1988). More recently, theoretical and 
empirical tests of this hypothesis have led to the asser- 
tion that loss or injury of the tail may instead be related 
to predator inefficiency or alternative sources of mor- 
tality and that careful investigation must be undertaken 
to determine the mechanism behind observed injury 
frequencies (Schoener 1979; Medel et al. 1988). While 
such data must indeed be interpreted with caution, re- 
porting the frequency of tail injury in study populations 
can be useful when combined with demographic data 
(Arnold 1988), for example in forming hypotheses re- 
garding sex or size-based differences in anti-predator 
mechanisms (Fitch 2003). 

Several researchers have reported the frequency of 
tail injuries in populations of snakes. Fitch (1999) found 
that the frequency of tail breakage increased with the 
age/size of individuals in a Kansas population of Nero- 
dia s. sipedon, the Northern Watersnake. More specif- 


ically, the frequency of female tail breakage ranged 
from 21.2 to 44.5%, and the frequency of male tail 
breakage ranged from 6.5 to 25% in increasing body 
size categories. Willis et al. (1982) reported that fe- 
male Eastern Garter Snakes (Thamnophis sirtalis) and 
Northern Ribbon Snakes (Thamnophis sauritus) had a 
higher incidence of tail loss than males (13% versus 
6%, and 12% versus 7%, respectively) and that tail loss 
was more prevalent in larger size classes. However, nei- 
ther trend was statistically significant in populations 
of Butler’s Garter Snake (Thamnophis butleri). Fitch 
(2003) found that tail breakage was more common for 
female T. sirtalis (16.7%) than for males (10.3%) and 
more common for large snakes (13.7%) than for first- 
year snakes (2.42%). In general, females and larger 
snakes appear more likely to have broken tails. 

I studied the frequency of broken tails of the North- 
ern Watersnake in the Beaver Archipelago of north- 
eastern Lake Michigan during the summers of 2000 
and 2001. I captured snakes from Beaver Island 
(45°41.26'N, 85°30.34'W), Garden Island (45°48.28'N, 
85°29.41'W), High Island (45°43.88'N, 85°39.54'W), 
and Hog Island (45°48.39'N, 85°22.15'W). Upon cap- 


436 


THE CANADIAN FIELD-NATURALIST 


TABLE 1. Chi-square contingency table for comparison of the frequency of tail breakage among male, female, and first-year 
Nerodia sipedon sipedon in the Beaver Archipelago of Lake Michigan. The calculated x? was 0.23 (critical x7 = 5.99, df =2). 


Class Number of Number of 

injured snakes uninjured snakes Total % Injured 
Male 7 55 62 AES 
Female 13 120 133 9.8 
First-year 2, 23 25 8.0 
Totals 22 198 220 10.0 


ture, I brought snakes to the laboratory where I then 
measured snout-vent length (SVL) and noted the tail 
condition and sex of each individual. I released all 
snakes at the point of capture as soon as possible 
after processing. I pooled snakes from all islands and 
calculated the overall percentage of snakes with tail 
breaks. I also compared the frequency of broken tails 
among males, females, and unsexed first-year snakes, 
and among three capture locations (Beaver Island Lake 
Michigan shoreline, Beaver Island interior lake, and 
Garden Island Lake Michigan shoreline) within sexes 
using Chi-square contingency table analysis. I hypoth- 
esized that females would have a higher frequency of 
tail breakage than males and that first-year snakes 
would have a lower frequency of tail breakage than 
both males and females. 

Two hundred and twenty snakes were captured from 
the four islands. Beaver Island produced 153 snakes, 
Garden Island 55, High Island 1, and Hog Island 11 
snakes. Captured snakes ranged in size from 149 to 
860 mm SVL with a mean of 478.2 mm and a stan- 
dard deviation of 164 mm. In all, 10% (22) of the 
snakes had broken tails. There was no statistically sig- 
nificant difference in the frequency of tail breakage 
among males, females, and first-year snakes (x7 = 0.23, 
critical y? = 5.99, df = 2; Table 1). Within males, there 
was no statistically significant difference in the fre- 


quency of tail breakage among capture sites (x? = 0.33, 
critical y7 = 5.99, df =2; Table 2). However, females 
captured on the Lake Michigan shoreline of Garden 
Island had a higher frequency of tail breakage than 
those captured elsewhere (x7 = 8.9, critical y? = 5.99, 
dfi=2; Table3): 

My data do not closely match those of any pre- 
vious studies. The overall frequency of tail breakage is 
similar to that reported by Willis et al. (1982). How- 
ever, unlike the populations of N. s. sipedon and T. 
sirtalis studied by Fitch (1999; 2003), and the popu- 
lations of T. sirtalis and T. sauritus studied by Willis 
et al. (1982), the frequency of broken tails is not great- 
er for females in the Beaver Archipelago overall (al- 
though it is greater if only Garden Island is consid- 
ered). Furthermore, first-year snakes in the Beaver 
Archipelago do not appear to have a lower frequency 
of tail breakage than larger snakes. The 7. butleri pop- 
ulations of Willis et al. (1982) appear to be similar to 
my snakes in the lack of common patterns of tail break- 
age. The sample size (287) for T. butleri in that study 
is similar to my own, while the sample sizes of the 7. 
sirtalis, T. sauritus and N. s. sipedon studies are con- 
siderably larger. This suggests that the ability to detect 
patterns in the frequency of tail injuries in snake pop- 
ulations may be dependent upon sample size. 


TABLE 2. Chi-square contingency table for comparison of the frequency of tail breakage of male Nerodia sipedon sipedon 
among capture sites in the Beaver Archipelago of Lake Michigan. The calculated x? was 0.33 (critical y° = 5.99, df =2). 


Capture Number of 
site 

Beaver Island Lake Michigan shoreline 
Beaver Island interior lakes 


Garden Island Lake Michigan shoreline 
Totals 


mee 


injured snakes 


Number of 
uninjured snakes Total % Injured 
ils) 17 11.8 
20 22 9.1 
18 21 14.3 
53 60 Hike 7/ 


TABLE 3. Chi-square contingency table for comparison of the frequency of tail breakage of female Nerodia sipedon sipedon 
among capture sites in the Beaver Archipelago of Lake Michigan. The calculated x7 was 8.9 (critical y* = 5.99, df =2). 


Capture site Number of 


injured snakes 


Beaver Island Lake Michigan shoreline 
Beaver Island interior lakes 
Garden Island Lake Michigan shoreline 


Totals 


= 
wlnony sp 


Number of 
uninjured snakes Total % Injured 
60 64 6.2 
34 36 SD) 
21 28 25.0 
TS 128 10.1 


2004 


Fitch (2003) suggested that the frequency of tail 
breakage should be expected to vary among different 
study sites based on the presence of predators and 
availability of escape cover. Such factors may explain 
the differences in overall frequency of tail breakage 
between this and other studies and between islands in 
the Beaver Archipelago within females. Fitch (1999) 
also suggested that NV. s. sipedon individuals with great- 
er mass are more likely to have broken tails because 
they whirl repeatedly in an attempt to escape when 
grasped by a predator. Greater mass makes tail break- 
age more likely when this whirling motion is employed. 
The smaller mass of first-year snakes, the smaller mass 
of male N. s. sipedon in comparison to females of the 
same age (Fitch 1999), and the fact that broken tails 
may potentially damage the hemipenes of males all sug- 
gest that these groups should have lower frequencies 
of tail breakage than large snakes and females, res- 
pectively, regardless of locality. These patterns were 
not clear in this study or in the populations of 7: 
butleri studied by Willis et al. (1982). 

Sample size may be the critical factor in the dis- 
crepancy between studies. The two studies that did 
not show common patterns in tail breakage had rela- 
tively small sample sizes. A large sample size may be 
necessary for two reasons: (1) the patterns are real 
but the categorical nature of the data (break or no 
break) requires that large samples be collected for a 
pattern to be detectable, or (2) the patterns are not 
real but stem from the fact that any difference, no 
matter how small, can be statistically significant if 
the sample size is large enough (Johnson 1999). The 
general similarity of patterns of tail breakage across 
studies with large sample sizes suggests that the first 
reason may be the case. However, discrepancies among 
studies underscore the need for more research in this 
area. Researchers should continue to record and 
report the frequency of tail breakage in populations 
of lizards and snakes, but more carefully designed 
studies that attempt to determine the direct causes 
and consequences of tail breakage are needed (Medel 
et al. 1988). Until such studies are carried out, the 
meaning and utility of tail breakage data will remain 
speculative. 


NOTES 


437 


Acknowledgements 

This research was done as part of a M.S. thesis at 
Central Michigan University. I thank my Thesis com- 
mittee: James C. Gillingham, Michael J. Hamas, and 
David L. Clark. I also thank John W. Rowe for help 
and advice in the field and laboratory. E. S. Bowen, 
M. F. Haussmann, and two anonymous reviewers pro- 
vided valuable comments on earlier drafts. Financial 
support was provided by the Central Michigan Uni- 
versity Department of Biology, the Central Michigan 
University College of Graduate Studies, and the 
Central Michigan University Biological Station on 
Beaver Island. Animals were collected under MDNR 
Cultural and Scientific Collector’s Permit CA 341 is- 
sued to J. C. Gillingham. The experimental protocol 
was approved by the Central Michigan University In- 
stitutional Animal Use and Care Committee (approval 
number 02-06). 


Literature Cited 

Arnold, E. N. 1988. Caudal autotomy as a defense. Pages 235- 
273 in Biology of the Reptilia. Volume 16, Ecology B. Edited 
by C. Gans and R. B. Huey. Alan R. Liss Inc., New York. 

Fitch, H. S. 1999. A Kansas snake community: composition 
and changes over 50 years. Krieger Publishing Company, 
Malabar, Florida xi + 165 pages. 

Fitch, H. S. 2003. Tail loss in Garter Snakes. Herpetological 
Review 34: 212-213. 

Johnson, D. H. 1999. The insignificance of statistical signifi- 
cance testing. Journal of Wildife Management 63: 763- 
TLE: 

Medel, R. G., J. E. Jimenez, S. F. Fox, and F. M. Jaksic. 
1988. Experimental evidence that high population fre- 
quencies of lizard tail autotomy indicate inefficient preda- 
tion. Oikos 53: 321-324. 

Mendelson, J. R. 1992. Frequency of tail breakage in Conio- 
phanes fissidens (Serpentes: Colubridae). Herpetologica 
48: 448-455. 

Schoener, T. W. 1979. Inferring the properties of predation 
and other injury-producing agents from injury frequencies. 
Ecology 60: 1110-1115. 

Willis, L., S. T. Threlkeld, and C. C. Carpenter. 1982. Tail 
loss patterns in Thamnophis (Reptilia: Colubridae) and the 
probable fate of injured individuals. Copeia 1982: 98-101. 


Received 21 February 2003 
Accepted 4 November 2004 


438 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


The Heather Vole, Genus Phenacomys, in Alaska 


S. O. MACDoNALD!:3, AMY M. RUNCK?, and JosEPH A. COOK>*” 


1P.O. Box 58, Gila, New Mexico 88038-0058 USA 


Biological Sciences Department, Idaho State University, Pocatello, Idaho 83209-8007 USA 
3Museum of Southwestern Biology, University of New Mexico, Albuquerque, New Mexico 87131 USA; Corresponding 


author, e-mail cookjose @unm.edu 


MacDonald, S. O., Amy M. Runck, and Joseph A. Cook. 2004, The heather vole genus Phenacomys, in Alaska. Canadian 


Field-Naturalist 118(3): 438-440. 


Four specimens of heather vole (genus Phenacomys) collected in the coastal mountains of Southeast Alaska document the 
first Recent records of this vole in Alaska. Alternative hypotheses on the relationship of these newly-discovered populations 
to extant and historical populations are outlined, and additional studies proposed. 


Key Words: Heather vole, Phenacomys sp., Southeast Alaska. 


Since publication of a synopsis of the land mammal 
fauna of Southeast Alaska (MacDonald and Cook 
1996), new field and laboratory studies of the mam- 
mals of this region have resulted in new perspectives 
on the zoogeography of the North Pacific Coast (Con- 
roy et al. 1999; Cook and MacDonald 2001; Cook et 
al. 2001). In our 1996 publication, we speculated that 
several mammal species found on the eastern side of 
the coastal mountains in Canada may occur along the 
Alaska coast. One species, the heather vole (Phena- 
comys intermedius), had previously been recorded 
within 2 km of the southern Alaska border in the sub- 
alpine near Salmon Glacier in British Columbia (ap- 
proximately 56.17° N, 130.03° W), and Nagorsen and 
Jones (1981) reported its occurrence in the Chilkat Pass 
north of Haines and Southeast Alaska. Subsequent 
trapping efforts in Alaskan mountains near Salmon 
Glacier and elsewhere along the mainland coast result- 
ed in the capture of four specimens of this vole (Table 
1) from two widely separate localities (Figure 1). 
These specimens, deposited at the University of Alaska 
Museum (UAM), constitute the first documented rec- 
ords of extant populations of heather vole in Alaska. 

On 11 August 1996, we captured a single heather 
vole (UAM 42371) above timberline in the glacially- 
isolated Chilkat Range at the northern end of South- 
east Alaska near Excursion Inlet, Glacier Bay National 
Park (58.41° N, 135.43° W; elevation approximately 
900 m). Also recorded in this alpine area were Sorex 
monticolus and melanistic Marmota caligata. 

In 1999, three heather voles were collected from 
the far southern end of mainland Southeast Alaska, 
approximately 400 km south of the 1996 Glacier Bay 
record. On 23 September 1999, two male (UAM 64239, 
UAM 64240) and one female (UAM 50000) heather 
voles were found in subalpine habitat along the Titan 
Trail in the Reverdy Mountains, approximately 10 
km north of Hyder, Alaska (approximately 55.83° N, 
130.03° W; elevations between 1128-1220 m). Synap- 
tomys borealis was also sampled at this site. 


The distribution of the heather vole is poorly known 
throughout western North America (Figure 1) and geo- 
graphic variation has been little studied (for overviews 
see McAllister and Hoffmann 1988; Nagorsen 1990; 
Musser and Carleton 1993). Some authorities (Ander- 
son 1942; Cowan and Guiguet 1965; Peterson 1966; 
Hallett 1999) recognized the distinctiveness of western 
populations (P. intermedius of westcentral British 
Columbia south to California and New Mexico) and 
separated them from P. ungava in the north and east 
(southern Yukon Territory and northern British Colum- 
bia eastward to Newfoundland) (McAllister and Hoff- 
mann 1988; George 1999); others considered all to 
comprise a single species (Crowe 1943; Youngman 
1975; Honacki et al. 1982; Nagorsen 2004). Hall (1981) 
speculated that intermedius may be a composite of two 
or three allopatric species (see McAllister and Hoff- 
mann 1988), but the lack of specimens from too few 
areas has prevented meaningful progress in resolving 
the problem. Reluctance of this species to enter traps 
may be the primary factor limiting sample size and 
study materials (McAllister and Hoffmann 1988). 

The taxonomic affinities of newly-discovered popu- 
lations of heather vole in Southeast Alaska are not 
immediately evident. Based on the map provided by 
Hall (1981: 787, as adjusted by McAllister and Hoff- 
mann 1988 and Nagorsen 2004), Alaska’s southern- 
most heather voles could share close kinship with the 
P. i. intermedius group of “western” heather voles 
occurring in the coastal mountain ranges of adjacent 
British Columbia. Heather voles from northern South- 
east Alaska, on the other hand, might possibly have 
connection with the “eastern” populations (as P. unga- 
va mackenzii) in northwestern British Columbia and 
southern Yukon Territory. Both of these scenarios pre- 
sume relatively recent colonization from inland pop- 
ulations following the (ongoing) retreat of the last 
major glaciation from the region. A similar scenario 
has been proposed for American Marten (Martes amer- 
icana) (Stone et al. 2002) and other mammals (Cook 


438 


2004 NOTES 439 


TABLE |. Summary of specimen data from collection notes and records at the University of Alaska Museum (UAM). Alaska 
Frozen Tissue Number refers to the cryogenic collection administered by UAM. Measurements and abbreviations are as 
follows: total length-tail length-hind foot-ear from notch (in millimeters)=mass (in grams); t = testes (length and width in 
mm), plsc = number of placental scars on right and left uterine horn; F = female, M = male. 


Alaska Frozen Tissue Number 


UAM Catalog Number 


Sex (Reproduction) Measurements 


42371 17262 
50000 30176 
64239 30178 
64240 30179 


et al. 2001). An alternative hypothesis, however, 1s that 
one or more of these populations are remnants of a 
former coastal population whose origin in the region 
preceded the last glacial advance (MacDonald and 
Cook 1996; Fleming and Cook 2002). 

The discovery of late Pleistocene fossil remains of 
Phenacomys in limestone cave deposits in the Alex- 
ander Archipelago of Southeast Alaska has added his- 
torical dimension to these hypotheses. Preliminary 


Chilkat River 


Gulf 
of Alaska 


LOCALITY KEY 
@ Chilkat Range 
@ Reverdy Mts. 


On Your Knees 
Cave fossil site 


M (t=2X3) 120-31-19-11=19 

F (plsc: 4R, OL) 156-37-18-14=33.9 
M (t=3X2) 129-32-17-13=22.8 
M (t=3X2) 115-26-16-10=14.6 


analyses of thousands of rodent teeth recovered from 
On Your Knees Cave at the extreme northwest corner 
of Prince of Wales Island in Southeast Alaska (Figure 
1) (Heaton 1995, 2004*; Heaton and Grady 2003) in- 
dicate that Heather Voles had been present in the vicin- 
ity of this site during the Middle Wisconsin up to the 
start of, but not during or following, the last glacial 
maximum (between about 24 000 to 13 000 years 
ago) (Heaton 2004*; Heaton and Grady 2003). Other 


% CANADA 


Sc 

, 
Za 
4p 


Stikine River 


Unuk River 


Ficure 1: Localities of specimens of Phenacomys in Southeastern Alaska from Recent and Late Pleistocene sites. Inset range 
map modified according to McAllister and Hoffman (1988) following subspecific designations suggested by George 


(1999) and Hallett (1999). 


440 


rodent remains discovered in these same pre-glacial 
deposits were Brown Lemming, Lemmus trimucro- 
natus, a species thought absent from the region but 
recently discovered by UAM in the mountains near 
Haines, and the Hoary Marmot, Marmota caligata, a 
species now restricted to mainland localities. 

How the new records of Phenacomys from South- 
east Alaska relate to extant and historical populations 
has yet to be determined. Analyses of samples using 
modern molecular and morphological techniques could 
help clarify these relationships and provide a test of 
alternate hypotheses on the origin and dynamics of 
the region’s biota. Clearly, more samples from more 
localities are needed. An expanded effort to excavate 
fossils throughout the region, along with a continua- 
tion of survey efforts to further define the current 
distribution of Phenacomys and other mammals that 
share a deep history in the region (e.g., Lemmus, 
Marmota), as well as provide adequate samples for 
assessment, are needed. The alpine zone of Southeast 
Alaska is virtually unexplored and future inventories 
need to target sites above treeline along the mainland 
as well as on the more mountainous islands in the 
Alexander Archipelago. 


Acknowledgments 

A number of individuals assisted with field work 
that led to these new records including C. T. Seaton, 
Able Duffy, Joni Reese, Mike Brown, Guillermo D’Eha, 
and John Chythlook. Brandy Jacobsen, Gordon Jarrell, 
and Dusty McDonald of the University of Alaska Mu- 
seum provided specimen information. Thanks also to 
David Nagorsen for reviewing an early draft of this 
paper. This research was funded by the United States 
Fish and Wildlife Service, United States Department of 
Agriculture Forest Service, Alaska Cooperative Fish 
and Wildlife Research Unit, and the National Science 
Foundation (DEB0196095 & DEB9981915). 


Documents Cited [marked * in text citations] 

Heaton, T. H. 2004. Ice Age paleontology of Southeast Al- 
aska. http://www.usd.edu/esci/alaska/index.html. 3 Decem- 
ber 2004. 


Literature Cited 

Anderson, R. M. 1942. Canadian voles of the genus Phena- 
comys with description of two new Canadian subspecies. 
Canadian Field-Naturalist 56: 56-60. 

Conroy, C. J., J. R. Demboski, and J. A. Cook. 1999. 
Mammalian biogeography of the Alexander Archipelago 
of Alaska: a north temperate nested fauna. Journal of 
Biogeography 26: 343-352. 

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Cook, J. A., A. L. Bidlack, C. J. Conroy, J. R. Demboski, 
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MacDonald. 2001. A phylogeographic perspective on 
endemism in the Alexander Archipelago of southeast 
Alaska. Biological Conservation 97: 215-227. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Cowan, I. M., and C. J. Guiguet. 1965. The mammals of 
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fauna of southeast Alaska. Canadian Field-Naturalist 
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nomic and geographic reference. Edited by D. E. Wilson 
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Received 26 November 2002 
Accepted 30 December 2004 


2004 NOTES 44] 


Extension de l’aire de distribution connue de la Musaraigne fuligineuse, 
Sorex fumeus, dans le nord-est du Québec 


JEAN-FRANCOIS DESROCHES! et ISABELLE PICARD 


' Collége de Sherbrooke, Département des Techniques d’écologie appliquée, 475 du Parc, Sherbrooke, Québec JIK 4K] 
Canada 


Desroches, Jean-Francois, et Isabelle Picard. 2004. Extension de l’aire de distribution connue de Ja Musaraigne fuligineuse, 
Sorex fumeus, dans le nord-est du Québec. Canadian Field-Naturalist 118(3) : 441-442. 


La Musaraigne fuligineuse (Sorex fumeus) figure sur la liste des especes de la faune vertébrée susceptibles d’étre désignées 
menacées ou vulnérables au Québec. Le 7 juin 1999, nous avons récolté un individu mort en bordure de la riviére Saint-Nicolas, 
a Godbout, dans la région de la Céte Nord. Cette observation constitue une extension de l’aire de répartition connue de |’ espéce 


d’ environ 200 km vers le nord-est. 


Mots-clés : Musaraigne fuligineuse, Sorex fumeus, aire de distribution, Québec. 


La Musaraigne fuligineuse (Sorex fumeus) est pré- 
sente dans le nord-est de 1’ Amérique du Nord, des 
Great Smokies des Carolines au sud jusqu’au centre 
de l’Ontario et aux Maritimes vers le nord (Banfield 
1977). Au Québec, la mention la plus nordique de 
Pespéce se situe a Val-Jalbert, aux abords du lac Saint- 
Jean (van Zyll de Jong 1983). Sa répartition connue vers 
le nord-est, sur la rive nord du fleuve Saint-Laurent, ne 
dépasse guére la riviére Saguenay (van Zyll de Jong 
1983; Banfield 1977). Cette musaraigne figure sur la 
liste des espéces de vertébrés susceptibles d’étre désig- 
nées menacées ou vulnérables au Québec (Beaulieu 
1992). Les chercheurs contactés en 1989 n’avaient pas 
rapporté de capture récente au Québec (Beaulieu 1992) 
mais depuis, elle a été capturée a divers endroits notam- 
ment dans le sud de la province (G. Lupien, commu- 
nication personnelle). 

Le 7 juin 1999, nous avons trouvé un spécimen de la 
Musaraigne fuligineuse mort, probablement noyé, en 
bordure de la riviére Saint-Nicolas, a environ 100 métres 
au sud de la route 138 a Godbout (49°19'10"N, 
67°47'24"0), dans la région de la Cote Nord. Lhabitat 
a cet endroit est une aulnaie, bordée par une sapiniere 
a Bouleau blanc. Aucun autre micromammifere n’y a 


été observé. Le spécimen de Musaraigne fuligineuse 
a été récolté pour identification. Les mesures ont été 
prises sur le spécimen a l’aide d’un vernier a cou- 
lisse, par J-F. D. et S. Houde, et elles correspondent a 
celles de la Musaraigne fuligineuse (tableau 1). Au 
Québec, la Musaraigne fuligineuse peut étre con- 
fondue avec la Musaraigne cendrée (Sorex cinereus), 
ayant toutes deux de quatre a cing dents unicuspides 
visibles, la limite postérieure du foramen infraorbi- 
taire antérieure ou prés de la limite entre les molaires 
1 et 2, le trou mentonnier situé a la hauteur du proto- 
conide de la moiaire 1, et la mandibule dépourvue de 
foramen postmandibulaire (Maisonneuve et al. 1997). 
Elle s’en distingue toutefois par certaines mesures du 
crane et du corps (tableau 1). Le spécimen a malheu- 
reusement été perdu quelques jours plus tard, jeté par 
erreur avec d’autres micromammiferes morts, mais les 
mesures prises confirment son identification. 

Cette observation constitue une extension d’aire de — 
Vespece de 200 km vers le nord-est, soit a partir de 
V’embouchure de la riviére Saguenay (van Zyll de Jong 
1983). La Musaraigne cendrée (Sorex cinereus), une 
espece plus commune au Québec (observation per- 
sonnelle), a une distribution qui s’étend bien plus au 


TABLEAU |. Mesures comparées du spécimen de musaraigne trouvé sur la Cote Nord, de la Musaraigne fuligineuse et de la 


Musaraigne cendrée. 


Longueur Largeur du Longueur Longueur de 
Espéce du crane (mm) maxillaire (mm) totale (mm) la queue (mm) 
Spécimen de la Céte Nord WET 4.8 111,4 47,8 
Musaraigne fuligineuse 17,5 a 18,8? 4,8 45,4? 1104127! 40 452! 
SIS? >4,6° 104 a 1252 42 a 54? 
Musaraigne cendrée 15,2 a 16,92 3.8 44,32 924110! 37 a 46! 
<173 <4,6° Ta le52 28 a 507 


' Banfield 1977 
> van Zyll de Jong 1983 
3 Maisonneuve et al. 1997 


441 


442 


nord (van Zyll de Jong 1983; Banfield 1977). La dé- 
couverte de la Musaraigne fuligineuse 4 Godbout in- 
dique que la distribution de certaines espéces de petits 
mammiféres est encore mal connue. En ce sens, il est 
permis de croire que de futures découvertes seront 
faites et permettront de mieux connaitre les habitats 
fréquentés par ces espéces et de préciser leur aire de 
distribution dans le nord de la province. II est a noter 
que cette mention de la Musaraigne fuligineuse figure 
dans |’Atlas des micromammiféres du Québec, sans 
documentation publié en 2002 (Desrosiers et al. 2002). 


Remerciements 

Nous tenons a remercier Mme Stéphanie Houde, 
technicienne de la faune, pour la vérification de l’iden- 
tification du spécimen de méme que M. Gilles Lupien, 
technicien de la faune a la Société de la Faune et des 
Parcs du Québec, pour ses commentaires sur les cap- 
tures de l’espéce au Québec. Nous remercions égale- 
ment le comité ZIP de l’estuaire du Saint-Laurent, qui a 
permis la réalisation de |’inventaire duquel la présente 
découverte de la Musaraigne fuligineuse a été faite. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Littérature citée 

Banfield, A. W. F. 1977. Les Mammiféres du Canada. Les 
Presses de l’ Université Laval et University of Toronto Press. 
Publié par le Musée national des Sciences naturelles, 
Musées nationaux du Canada. Ottawa, Canada. 406 pages. 

Beaulieu, H. 1992. Liste des espéces de la faune vertébrée 
susceptibles d’étre désignées menacées ou vulnérables au 
Québec. Ministére du Loisir, de la Chasse et de la Péche. 
Québec. 107 pages. 

Desrosiers, N., R. Morin, et J. Jutras. 2002. Atlas des 
micromammiféres du Québec. Société de la faune et des 
parcs du Québec et Fondation de la Faune du Québec. 92 
pages. 

Maisonneuve, C., R. Mc Nicoll, S. St-Onge, et A. Des- 
rosiers. 1997. Clé d’identification des micromammiferes 
du Québec. Ministére de 1’ Environnement et de la Faune, 
Québec. 19 pages. 

van Zyll de Jong, C. G. 1983. Traité des Mammiféres du 
Canada, volume | : Les Marsupiaux et les Insectivores. Mu- 
sée national des Sciences naturelles, Musées nationaux du 
Canada. Ottawa, Canada. 217 pages. 


Recu 28 octobre 2001 
Acceptée 15 janvier 2005 


Gulls, Larus spp., Foraging at Pink Salmon, Oncorhynchus gorbuscha, 


Spawning Runs 


Mary FE. WILLSON 


5230 Terrace Place, Juneau, Alaska 99801 USA; e-mail mwillson@ gci.net 
Willson, Mary F. 2004. Gulls, (Larus spp.), foraging at Pink Salmon, Oncorhynchus gorbuscha, spawning runs. Canadian 


Field-Naturalist 118(3): 442-443. 


Small and immature gulls foraged more often on drifting salmon eggs than did large and mature gulls, and large and mature 
gulls foraged more often on salmon carcasses, at streams in Southeast Alaska. These differences may be related to body size 
via physical strength and dominance status, as well as foraging experience. 


Key Words: Gulls, Larus spp., Pink Salmon, Oncorhynchus gorbuscha, Southeast Alaska. 


Salmon spawning runs attract many foraging birds 
and mammals (Willson and Halupka 1995) and are 
clearly an important food resource for wildlife in late 
summer and fall. Gulls (Larus spp.) are among the most 
numerous predators and scavengers at salmon runs, 
sometimes occurring in the thousands (Isleib and Kes- 
sel 1973). Here I report how four species of gull ex- 
ploited runs of Pink Salmon (Oncorhynchus gor- 
buscha) at the mouths of streams in Juneau, Alaska. 


Study Area and Methods 

Most of these observations were made at the mouths 
of Salmon Creek and Sheep Creek (less than 12 km 
apart), with some supplementary observations at four 
other small, nearby streams, within the city and bor- 
ough of Juneau in Southeast Alaska (ca. 58°30'N, 
133°30'W). I observed four species of gulls foraging 
at or near low tides, usually in the morning, from 27 
August to 6 September 2002 (>6 hrs of observation). 
At any one time, many dozens of gulls were present 
at the two primary study locations. At low tide, these 


creeks flow over intertidal deltas, often with several 
shallow channels used by incoming salmon. I scanned 
these deltas with spotting scope and binoculars, noting 
the foraging activities of the gulls and counting the 
individuals engaged in several distinctive foraging ac- 
tivities. Birds shifted position frequently, using different 
parts of the deltas; many individuals of the three larger 
species also changed foraging activity as I watched. 
Most scans were made at different locations on differ- 
ent days; if more than one scan occurred on the same 
day and stream delta, they were separated in time by 
about 30 min. I recorded foraging observations in three 
categories: foraging on salmon eggs (drifting eggs are 
common because later spawners often disturb the grav- 
els over the nests of earlier spawners, foot-paddling 
by gulls also stirs up poorly buried eggs), foraging on 
salmon carcasses (including some extraction of eggs 
from living or dead fish), and searching for inverte- 
brates in Fucus mats (where amphipods and isopods 
were common) and tide pools. 


2004 NoTes 443 
TABLE |. Foraging activity of four species of gull at salmon spawning runs in southeast Alaska. 

Species/age class Eggs Salmon Invertebrates N (observations) 
Bonaparte’s Gull 82% - 18% 353 

Mew Gull adults 33% 1% 66% 232 

Herring Gull adults* 31% 58% 11% 26 

Herring Gull immatures 80% 20% - 15 
Glaucous-wing adults* 11% 87% 2% 362 
Glaucous-wing immatures 44% 55% 1% 222 


‘Frequency distributions for adults and immatures were significantly different (y’, = 88.5 and 7.1, for the two species 


respectively; p < 0.05 in both cases). 


Results 

Foraging behavior of each species was very con- 
sistent among the observed streams, but there were 
marked differences in foraging among species and 
age classes (Table 1). The small (ca. 190 g, Sibley 
2000) Bonaparte’s Gull (L. philadelphia) foraged for 
salmon eggs chiefly by hovering over the streams and 
plunge-diving for drifting eggs, but they occasionally 
sought invertebrates in Fucus mats or tide pools. 
Most birds were in nonbreeding plumage, and adults 
were not distinguished from immatures. 

Mew Gulls (L. canus), of intermediate size (ca. 
420 g, Sibley 2000), waded in the shallows, searching 
for drift eggs, or hunted invertebrates in Fucus mats. 
They seldom foraged on fish carcasses and then only 
if no large gulls were nearby. Too few immatures were 
present for adequate sampling, so all observations per- 
tain to adults. 

Both Herring (L. argentatus) and Glaucous-winged 
(L. glaucescens) gulls (body mass about 1000-1100 g 
for both species, Sibley 2000) foraged chiefly on car- 
casses and on eggs, only rarely visiting the Fucus mats 
on these deltas. Eggs were commonly obtained while 
wading, occasionally by foot-paddling. Glaucous-wing 
adults sometimes pulled live salmon from the stream, 
poking initially at eyes and at the vent area to force egg 
extrusion. Immatures of both species foraged more 
often on eggs than did adults, while adults foraged more 
often on carcasses. Both of these larger gulls often 
examined very old, bleached and decayed, carcasses 
that appeared to offer little or nothing edible. 


Discussion 

Salmon eggs are rich in lipids (Gende et al. 2001, 
2004), offering an easily captured and high-calorie 
food, and brightly colored, so they are easy to see as 
they drift downstream. Bonaparte’s Gulls, Mew Gulls, 
and immatures of the larger species foraged heavily 
on eggs. The demands of imminent fall migration for 
the smaller gulls may provide added impetus for 
rapid energy acquisition; most Bonaparte’s Gulls left 
the area before the Pink Salmon runs were completely 
over. The immatures are less experienced foragers than 
adults, and eggs may be more easily accessed than 


carcasses. Carcasses are probably less accessible to 
smaller gulls because the skin is tough and hard to 
breach and also because the larger gulls (and ravens 
and eagles) can easily dominate the smaller birds. 
Glaucous-winged Gulls selectively killed and eviscer- 
ated female Sockeye Salmon (O. nerka) in shallow 
streams in westem Alaska (Mossman 1958). The forag- 
ing differences among gull species here are similar to 
the observations of Moyle (1966) elsewhere in Alaska. 

Other species of wildlife also favor salmon eggs. 
Bears (Ursus spp.) often feed selectively on eggs (and 
other high-lipid body parts) from live-caught fish 
(Gende et al. 2001). American Dipper (Cinclus mexi- 
canus) chicks, juveniles, and adults feed on drift eggs 
(Obermeyer et al. 1999; K. E. Obermeyer and M. F. 
Willson, unpublished observations). Foraging: on drift 
eggs by wildlife has no significant effect on salmon 
populations, because drift eggs do not survive outside 
the nesting gravels. 


Literature Cited 

Gende, S. M., T. P. Quinn, and M. F. Willson. 2001. Con- 
sumption choice by bears feeding on salmon. Oecologia 
127: 372-382. 

Gende, S. M., T. P. Quinn, M. F. Willson, R. Heintz, and 
T. M. Scott. 2004. Magnitude and fate of salmon derived 
nutrients and energy in a coastal system ecosystem. Journal 
of Freshwater Ecology 19: 149-160. 

Isleib, M. E., and B. Kessel. 1973. Birds of the north gulf 
coast-Prince William Sound region, Alaska. Biological 
Papers, University of Alaska 14: 1-149. 

Mossman, A. S. 1958. Selective predation of glaucous-winged 
gulls upon adult red salmon. Ecology 39: 482-486. 

Moyle, P. 1966. Feeding behavior of the glaucous-winged 
gull on an Alaska salmon stream. Wilson Bulletin 78: 
175-190. 

Obermeyer, K. E., A. Hodgson, and M. F. Willson. 1999. 
American Dipper, Cinclus mexicanus, foraging on salmon 
eggs. Canadian Field-Naturalist 113: 288-290. 

Sibley, D. A. 2000. The Sibley guide to birds. Knopf, New 
York. 

Willson, M. F., and K. C. Halupka. 1995. Anadromous fish 
as “keystone” resources in vertebrate communities. Con- 
servation Biology 9: 489-497. 


Received 26 February 2003 
Accepted 22 January 2004 


444 THE CANADIAN FIELD-NATURALIST Vol. 118 


Sequential Polyandry in Piping Plover, Charadrius melodus, Nesting 
in Eastern Canada 


DIANE L. AMIRAULT!, JONATHAN KIERSTEAD!, PETER MACDONALD? and LARRY MACDONNELL! 


! Canadian Wildlife Service, Environment Canada, P.O. Box 6227, Sackville, New Brunswick E4L 1G6 Canada 
? Nova Scotia Department of Natural Resources, P.O. Box 99, Tusket, Nova Scotia BOW 3MO Canada 


Amirault, Diane L., Jonathan Kierstead, Peter MacDonald and Larry MacDonnell. 2004. Sequential polyandry in Piping 
Plover, Charadrius melodus, nesting in eastern Canada. Canadian Field-Naturalist 118(3): 444-446. 


On Cape Sable Island, Nova Scotia, we confirmed that a banded female Piping Plover (Charadrius melodus melodus) produced 
two broods of chicks during the 2000 nesting season, the second on a beach approximately 2 km from the first. The female 


abandoned her second brood two days after hatching, leaving the male to complete brood rearing. 


Key Words: Piping Plover, Charadrius melodus, sequential polyandry, Nova Scotia. 


Piping Plovers (Charadrius melodus) generally have 
been thought to be monogamous, with males estab- 
lishing and defending territories and raising only one 
brood per season (Wilcox 1959; Cairns 1982; Haig 
and Oring 1988; Haig 1992). Renesting in Piping 
Plovers is common if a clutch is lost (Haig 1992) and 
occasionally occurs if a brood of chicks is lost (Mac- 
Ivor 1990). In Manitoba, Piping Plovers exhibited 
sequential polyandrous behavior (Oring 1982; Haig 
1992) where some breeding adults had multiple nests 
and mates (Haig and Oring 1988) after predation or 
storm events caused nest loss. 

The best example of polyandry in shorebirds has 
been documented in the Spotted Sandpiper (Actitis 
macularia) (Oring and Knudson 1973). Within Spotted 
Sandpipers, females are larger and more aggressive than 
males and establish and defend nesting territories. 
Female Spotted Sandpipers lay several clutches of eggs 
while different males incubate eggs and complete 
brood-rearing tasks for each. 

Bottitta et al. (1997) published the first observations 
of Piping Plovers successfully fledging two broods in 
one nesting season, at Griswold Point in Old Lyme, 
Connecticut, and Assateague Island National Seashore, 
along the Maryland and Virginia coast. During the 
Bottitta et al. (1997) studies (1989-1994), eight Piping 
Plover pairs successfully raised two broods each within 
a reproductive season. Although not all birds involved 
were banded to enable identification of individuals, 
mates in most cases were thought to have been re- 
tained because of plumage characteristics and proximity 
of second nests to locations of first nests. 

The potential for Piping Plovers P. m. meliodius to 
raise a second brood in eastern Canada is thought to 
be small, owing to the later, and therefore shorter, nest- 
ing season associated with a more northerly location. 
With aid of a banding program, we documented the 
production of a second brood by the same female 
Piping Plover in southwestern Nova Scotia during 
the 2000 nesting season. 


Study Area and Methods 

Piping Plovers have been closely monitored on two 
beaches (Daniels Head beach [43°26'N, 65°36'W] and 
Stoney Island beach [43°28'N, 65°34'W]) on Cape 
Sable Island in southern Nova Scotia since 1997 (P. 
MacDonald, unpublished data). Daniels Head beach 
encompasses a 15-90 m (high-low tide) wide stretch 
of light-coloured sand extending the full length of the 
3.3 km beach, broken by four rocky points and an inlet 
to a sandy tidal-flat area. Low vegetated (Ammophila 
breviligulata and other grasses) dunes provide a break 
between the beach and a brackish inland marsh. Sto- 
ney Island is a light-coloured sand beach bordered by 
low, densely vegetated dunes (Ammophila breviligulata, 
grasses, and shrubs). The 15-75 m (high-low tide) 
wide beach extends 1.8 km in a crescent shape from 
a rocky point in the south to an embankment in the 
north. The beaches are separated by an approximate- 
ly 2 km long coastal stretch characterized by rocky 
shoreline, a tidal channel, and headland on which a 
fish processing plant is located (Boates et al. 1994*). 
Daniel’s Head beach supported from three to seven 
nesting pairs of Piping Plovers in the 1997 to 2000 
period. Conversely, the single pair at Stoney Island 
beach was the only successful nesting event since 
LOOT: 

We trapped and banded birds on several occasions 
during May-July 2000. A modified Weller trap was 
used to capture adult birds incubating eggs (Weller 
1957). A trapped bird was immediately removed from 
the trap, banded with a United States Fish and Wildlife 
Service numbered metal incoloy band on the lower 
right leg (for adult birds) and a Canadian Wildlife 
Service bicoloured plastic band on the lower left leg 
(specific to province) and released. 


Results 

On 24 May 2000, we trapped and banded a female 
Piping Plover incubating four eggs on Daniels Head 
beach. Four chicks hatched from this clutch on 30 May. 
The female was not observed with the brood after 13 


2004 


June, at which time only one chick remained. This 
chick was considered fledged on 25 June. The adult 
male remained with the juvenile and both were ob- 
served together until 5 July. A nest containing three 
eggs was discovered on 23 June on Stoney Island 
beach. Four eggs were confirmed on 25 June. We 
trapped a female bird on this nest on 29 June and her 
leg band number confirmed that this individual was 
the female banded on 24 May at Daniels Head beach. 
This second nesting event appears to have been com- 
pleted with a different mate as the male from the first 
nest remained with the surviving chick until 5 July, 
well after fledging. The second pair hatched four eggs 
on 18 July and the female was not observed with the 
brood (4 chicks still remained) after 20 July. One 
chick from this brood was considered fledged on 13 
August, the last day both the juvenile and adult male 
were observed. Males from each nest were unbanded, 
facilitating identification of which adult (male or 
female) was present with the chicks. 


Discussion 

Although Haig and Oring (1988) documented poly- 
andry in Manitoba, their observations included renest- 
ing following nest loss but never more than one brood 
per nesting season. Our observations represent the first 
documented occurrence of a female Piping Plover 
relocating to produce a second brood within a nesting 
season on Canadian nesting grounds. Bottitta et al. 
(1997) were the first to report Piping Plovers raising 
more than one brood in a season. However, their oc- 
currences were documented in the United States and 
on a single nesting beach. Although closely related 
species such as Killdeer (Charadrius vociferus) and 
Snowy Plover (Charadrius alexandrinus) are known 
to produce two broods, this behaviour is thought to 
be extremely rare among Piping Plovers (Wilcox 1959; 
Cairns 1982; Whyte 1985; Haig and Oring 1988; 
Maclvor 1990; Strauss 1990; Loegering 1992). Thus, 
most second nest attempts are renests following nest 
loss. Presumably, the second nest on Stoney Island 
beach was initiated 18-19 June as Piping Plovers aver- 
age 6-8 days to complete laying of a full clutch (Haig 
1992). The interval between the last sighting of this 
female with her first brood (13 June) and the calcu- 
lated initiation of her new clutch is brief (5-6 days). 
However, Piping Plovers have been known to initiate 
renesting after a lost clutch in as few as 4 days 
(Whyte 1985; Haig and Oring 1988; MacIvor 1990; 
Loegering 1992). 

Bottitta et al.’s (1997) long-term study documented 
several examples of two broods being produced with- 
in a nesting season, all of which were thought to have 
been completed with the same mates. That is consis- 
tent with renesting Piping Plovers generally retaining 
the same mate (Wilcox 1959; Whyte 1985; Haig and 
Oring 1988; MaclIvor 1990). However, in southern 
Manitoba, renesting birds often chose a new mate in 


NOTES 


445 


cases of high predation and nest loss (Haig and Oring 
1988). Our record of intra-year mate switching sug- 
gests that switching may occur in cases where nest loss 
has not occurred and may increase an individual’s 
productivity. 

Several studies have indicated that it is not uncom- 
mon for one adult, usually the female, to abandon its 
brood prior to chicks fledging (Cairns 1977; Whyte 
1985; Haig 1987). However, this is the first docu- 
mented case of a female plover abandoning its brood 
in order to produce another nest. These observations 
indicate that a small number of females may leave 
early to renest and leave further parental care to the 
male. 

Previously documented cases of sequential poly- 
andry in Piping Plovers have been in circumstances 
associated with high predation rates and catastrophic 
storm events (Haig and Oring 1988; Maclvor 1990). 
Our observations demonstrate that sequential poly- 
andry may also occur where there is potential for 
increasing productivity. The documentation of a sec- 
ond brood produced by one female underscores the 
importance of having early nest attempts succeed. 
Such success would allow greater chance for a sec- 
ond nesting event to occur, resulting in greater prod- 
uctivity. Maclvor (1990) reported that more young 
fledge from first nest attempts than from renests and 
from early versus late nests. MacIvor (1990) suggest- 
ed that increased vulnerability of chicks later in the 
breeding season (July-August) was due to a greater 
human presence on beaches then. Haig and Oring 
(1988) reported greater parental care on first nest at- 
tempts versus renests; however, no difference in hatch- 
ing or fledging rates was noted. Therefore, measures to 
enhance nest success such as placing predator exclo- 
sures (Bottitta et al. 1997), symbolic fencing, and 
beach guardian patrols may be particularly important 
for first nest attempts as a successful early nest may 
increase reproductive success. 

The incidence of producing two broods within a nest- 
ing season is thought to be fairly low, but documenta- 
tion of one apparent double nesting event by an in- 
dividual has significant implications for conducting 
population counts of adult Piping Plovers. This event 
justifies adherence of any Piping Plover census to a 
brief window of time, as in the absence of informa- 
tion such as provided by this study, the second nesting 
event might have been erroneously recorded as two 
additional individuals. Counts within a prescribed time 
frame will decrease the potential for double-counting 
birds that have relocated to renest or produce a second 
brood. In past censuses, observers often have con- 
sidered an adult found on a nest or a single adult with 
a brood as a nesting pair (i.e., two adult birds), even 
if only one adult was observed. Our observations sug- 
gest that a small proportion of late-nesting pairs may 
actually be successful early nesters on a second nest 
attempt. Hence, danger in overestimating numbers 


446 


exists when counting adult Piping Plovers outside a 
census window as our observations indicate not all birds 
retain the same mate or beach in a nesting season. 


Acknowledgments 

The Canadian Wildlife Service funded the Piping 
Plover banding and monitoring program during the 
2000 season. The Nova Scotia Department of Natural 
Resources provided in-kind and logistical support for 
the 2000 seasonal effort. Andrew Boyne and Kevin 
Davidson provided useful comments on earlier drafts 
of the manuscript. 


Documents Cited (marked * in text) 

Boates, J. S., P. Austin-Smith, G. Dickie, R. Williams, and 
D. Sam. 1994. Nova Scotia Piping Plover Atlas. Unpub- 
lished Nova Scotia Department of Natural Resources report. 
86 pages. 


Literature Cited 

Bottitta, G. E., A. M. Cole, and B. Lapin. 1997. Piping Plov- 
ers produce two broods. Wilson Bulletin 109: 337-339. 

Cairns, W. E. 1977. Breeding biology of the piping plover 
in southern Nova Scotia. M.Sc. thesis, Dalhousie Univer- 
sity, Halifax. 115 pages. 

Cairns, W. E. 1982. Biology and behavior of breeding Pip- 
ing Plovers. Wilson Bulletin 94: 531-545. 

Haig, S. M. 1987. The population biology and life history 
patterns of the piping plover. Ph.D. dissertation, University 
of North Dakota, Grand Forks. 121 pages. 

Haig, S. M. 1992. Piping Plover. Pages 1-18 in The Birds 
of North America (2). Edited by A. Poole, P. Stettenheim, 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


and F. Gill. The Academy of Natural Sciences, Philadel- 
phia; The American Ornithologists’ Union, Washington, 
Die 

Haig, S. M., and L. W. Oring. 1988. Mate, site, and terri- 
tory fidelity in Piping Plovers. Auk 105: 268-277. 

Loegering, J. P. 1992. Piping Plover breeding biology, forag- 
ing ecology and behavior on Assateague Island National 
Seashore, Maryland. M.S. thesis, Virginia Polytechnic Insti- 
tute and State University, Blacksburg, Virginia. 

Maclvor, L. H. 1990. Population dynamics, breeding ecology, 
and management of Piping Plovers on outer Cape Cod, 
Massachusetts. M.S. thesis, University of Massachusetts, 
Amherst, Massachusetts. 

Oring, L. W. 1982. Avian mating systems. Pages 1-92 in 
Avian Biology. Volume VI. Edited by D. S. Farner and J. 
R. King. Academic Press, Inc., New York. 

Oring, L. W., and M. L. Knudson. 1973. Monogamy and 
polyandry in the spotted sandpiper. Living Bird 11: 59-73. 

Strauss, E. 1990. Reproductive success, life history patterns, 
and behavioral variation in a population of Piping Plover 
subjected to human disturbance. Ph.D. dissertation, Tufts 
University, Boston, Massachusetts. 

Weller, M. W. 1957. An automatic nest-trap for waterfowl. 
Journal of Wildlife Management 21: 456-458. 

Whyte, A. J. 1985. Breeding ecology of the Piping Plover 
(Charadrius melodus) in central Saskatchewan. M.Sc. 
thesis, University of Saskatchewan, Saskatoon, Saskatch- 
ewan. 

Wilcox, L. 1959. A twenty year banding study of the Piping 
Plover. Auk 76: 129-152. 


Received 23 May 2003 
Accepted 15 November 2004 


American Dipper, Cinclus mexicanus, Preys Upon Larval Tailed Frogs, 


Ascaphus truei 


Curisty A. MorRISSEY!:? AND ROBERTA J. OLENICK? 


'Department of Biological Sciences, Simon Fraser University, 8888 University Drive, Burnaby, British Columbia, V5A 1S6 
Canada 

>Never-Spook-the-Animals Wildlife Photography, 3778 West 13' Avenue, Vancouver, British Columbia, V6R 2S6 Canada 

>Corresponding author (current address): Canadian Wildlife Service, Environment Canada, 5421 Robertson Road, RR#1, 
Delta, British Columbia V4K 3N2 Canada; email: christy.morrissey @ec.gc.ca or cmorrissey @alumni.stu.ca 


Morrissey, Christy A., and Roberta J. Olenick. 2004. American Dipper, Cinclus mexicanus, preys upon larval tailed frogs, 
Ascaphus truei. Canadian Field-Naturalist 118(3): 446-448. 


The American Dipper (Cinclus mexicanus) is an aquatic songbird that inhabits fast-flowing mountain streams in western 
North America. Although dippers are known to feed primarily on aquatic invertebrates, they will also eat juvenile fish and 
salmon eggs when available. In 2002, while monitoring and photographing nesting activities of the American dipper, we 
observed and photographed adult dippers capturing Tailed Frog (Ascaphus truei) tadpoles and feeding them to their young. This 
note is intended to document a rarely observed occurrence and identify interactions between two relatively uncommon species. 


Key Words: American Dipper, Cinclus mexicanus, Tailed Frog, Ascaphus truei, feeding, Chilliwack River, British Columbia. 


American Dippers (Cinclus mexicanus) are North 
America’s only truly aquatic songbirds occupying 
mountain streams of western North America from 
Alaska south to Mexico (Kingery 1996). Dippers are 
known to feed almost exclusively on in stream fauna, 
diving underwater and probing among the rocks for 


benthic invertebrates and small fish (salmon and trout) 
as well as fish eggs (Kingery 1996). To our knowledge, 
there have been no previous records in the literature 
of dippers feeding on amphibians. However, while 
studying and photographing American Dippers in 
southwestern British Columbia, we observed dippers 


2004 


NOTES 


447 


ni eominstiscgagty em 


sa 


Creek, British Columbia, in June 2002. Photograph by Roberta Olenick. 


on several occasions feeding on Tailed Frog (Ascaphus 
truei) larvae. 

Field work on the American Dipper was conducted 
in the Chilliwack River watershed located in the Cas- 
cade Mountains of southwestern British Columbia, 
Canada. While observing dipper nests in June and July 
2002, we noted at least two different pairs of Ameri- 
_ can Dippers, attending separate nests on Tamihi Creek, 
_ a tributary of the Chilliwack River (49.02 N, 121.50 W, 


Ficure 2: Adult American Dipper delivering Tailed Frog tad- 
pole to newly fledged dipper on Tamihi Creek, British 
Columbia in July 2002. Photograph by Roberta Olenick. 


elevation 545 m) feeding Tailed Frog tadpoles to their 
young. Roberta Olenick photographed those dippers 
delivering Tailed Frog tadpoles to nestlings or fledg- 
lings at each nest site (Figures | and 2). 

On one occasion, an adult dipper captured a very 
large Tailed Frog tadpole (estimated length 5-6 cm) 
and attempted to feed it to one of its fledglings. The 
fledgling dipper struggled to swallow the large prey 
item without success. Several times, the adult dipper 
retrieved the tadpole, knocked it vigorously against a 
rock and manipulated the large wide head in its bill. 
Eventually, after several minutes of handling by the 
adult, the fledgling dipper managed to swallow the 
large tadpole whole. 

During routine annual benthic invertebrate collec- 
tions in April 1999-2002, we infrequently captured 
Tailed Frog tadpoles in our Surber samplers using 
kick-sampling procedures. Larval Tailed Frogs have 
a distinct round mouth, well adapted to use as suction 
onto the surface of in stream rocks, making them 
relatively easy to identify. Tailed Frog larvae were 
found on several tributaries of the Chilliwack River 
including Tamihi Creek, Borden Creek, and Chipmunk 
Creek. It is worth noting that all of these creeks have 
present or past logging activity which is thought to limit 
the presence and abundance of Tailed Frogs through 
increased water temperatures and siltation (Nussbaum 
et al. 1983). No other similar species of amphibians 
occupy fast flowing stream habitat in our study area. 

There are few known predators of Ascaphus truei. 
Tailed Frog tadpoles have been identified as impor- 


448 


tant prey for larval Pacific Giant (Dicamptodon ensa- 
tus) and Cope’s salamanders (Dicamptodon copei) and 
Red-legged Frogs (Rana aurora) (Bury 1968; Jones 
and Raphael 1998). Observations of garter snakes 
(Thamnophis spp.) (Karraker 2001) and Cutthroat 
Trout (Salmo clarki) depredating Tailed Frogs have 
also been previously reported (Daugherty and Shel- 
don 1982). We found only one instance in the litera- 
ture where American Dippers were cited as being a 
potential predator of Tailed Frogs, in Butler Creek, 
Montana (Daugherty and Sheldon 1982). However, it 
was not clear whether those authors actually observed 
dippers feeding on Tailed Frogs. Given that the Tailed 
Frog is a provincially blue-listed (vulnerable) species 
in British Columbia and of special concern in the na- 
tional COSEWIC listing, documentation of these rare 
observations is important. 

Tailed Frogs are endemic to the Pacific Northwest. 
They range from British Columbia south to Califor- 
nia, occupying western mountain streams and humid 
forests throughout a 15-20 year lifespan (Daugherty 
and Sheldon 1982). Larvae take from | to 4 years to 
metamorphose into adults in cool fast streams (Bull 
and Carter 1996). Their distribution directly overlaps 
that of the American Dipper, which occupies the same 
habitat and geographic range. Both species are consid- 
ered sensitive to environmental impacts in mountain- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


ous watersheds from anthropogenic sources (Nuss- 
baum et al. 1983, Kingery 1996). Given that these two 
species occupy the same habitat and geographic range, 
American Dipper predation of Tailed Frogs may be 
more widespread than previously acknowledged. 


Literature Cited 

Bull, E. L., and B. E. Carter. 1996. Tailed Frogs: Distri- 
bution, ecology, and association with timber harvest in 
northeastern Oregon. United States Forest Service Re- 
search Paper 497: 1-12. 

Bury, R. B. 1968. The distribution of Ascaphus truei in Cali- 
fornia. Herpetologica 24: 39-46. 

Daugherty, C. H., and A. L. Sheldon. 1982. Age determi- 
nation, growth, and life history of a Montana population 
of the tailed frog. Herpetologica 38: 461-468. 

Jones, L. and M. Raphael. 1998. Ascaphus truei (Tailed 
Frog) predation. Herpetological Review 29: 39. 

Karraker, N. E. 2001. Ascaphus truei predation. Herpeto- 
logical Review 32: 100. 

Kingery, H. E. 1996. American Dipper (Cinclus mexicanus). 
The birds of North America (229): 1-28. Edited by A. Poole 
and F. Gill. The Academy of Natural Sciences, Philadelphia, 
and American Ornithologists’ Union, Washington, D.C. 

Nussbaum, R. A., E. D. Brodie, and R. M. Storm. 1983. 
Reptiles and amphibians of the Pacific Northwest. Univer- 
sity Press of Idaho, Moscow, Idaho. 


Received 28 January 2003 
Accepted 6 December 2004 


Piping Plover, Charadrius melodus, egg viability after seawater immersion 


JULIE McKNiGut!, LINDA THOMAS?, and DIANE L. AMIRAULT? 


' Canadian Wildlife Service, Environment Canada, Atlantic Region, 45 Alderney Drive, Dartmouth, Nova Scotia B2Y 2N6 
Canada 

? Prince Edward Island National Park, Parks Canada, 2 Palmers Lane, Charlottetown, Prince Edward Island C1A 5V6 Canada 

3Canadian Wildlife Service, Environment Canada, Atlantic Region, 17 Waterfowl Lane, Sackville, New Brunswick E4L 1G6 
Canada 


McKnight, Julie, Linda Thomas and Diane L. Amirault. 2004. Piping Plover, Charadrius melodus, egg viability after seawater 
immersion. Canadian Field-Naturalist 118(3): 448-450. 


Four observed nest histories indicate Piping Plover eggs are able to survive immersion in seawater, but little is known 
regarding their hardiness. As Piping Plover nests are often exposed to tidal flooding, their eggs may be relatively resistant to 
inundation by seawater. Therefore, we suggest that replacing eggs recently flooded or washed out of-nests is a viable option 
for the recovery of individual nests. 


Key Words: Piping Plover, Charadrius melodus melodus, flooding, high tide, hatching success, viability, Prince Edward 


Island, Nova Scotia. 


The Atlantic Coast population of Piping Plover 
(Charadrius melodus melodus) is listed as Endan- 
gered in Canada and Threatened in the United States 
(Goossen et al. 2002: U.S. Fish and Wildlife Service 
1996). Nesting Atlantic coast Piping Plovers prefer 
flat coastal beaches with sand and pebble substrate 
(Boyne and Amirault 1999; Burger 1987; Cairns and 
McLaren 1980). The male scrapes a shallow depres- 
sion in the substrate between the mean high water mark 
and the edge of adjacent dunes or vegetation (Burger 


1987; Cairns 1982; Haig 1992). The young hatch 
after approximately 28 days and leave the nest within 
hours of hatching (Cairns 1982; Haig 1992). Nesting 
areas are often flooded by storm-induced overwashes 
(Cairns and McLaren 1980) and high water levels des- 
troy many plover nests each season (Sylvester 1991). 

Four cases in which flooded Piping Plover eggs 
remained viable were documented during the course 
of regular monitoring at Prince Edward Island National 
Park and in southern Nova Scotia. The following nest 


2004 


histories confirm the viability of flooded eggs. 


Case 1. On 29 May 1992, a nest at Cavendish Sand- 
spit in Prince Edward Island National Park (46.5036°N, 
63.4408°W) was completed with four eggs. On 5 June, 
the nest was submerged by a high tide. Both adults 
were present but not searching for their eggs or vocal- 
ising excessively. Their eggs were removed from the 
nest, temporarily replaced with clay eggs and returned 
to the nest later that day. The shells that had lined the 
nest were covered by wet sand and some water still 
remained in the nest. After a brief period, one of the 
adults returned to the nest and began to incubate and 
the second adult arrived at the site shortly thereafter. 
On 24 June, four chicks hatched after 26 days of 
incubation. The four chicks were considered to have 
fledged on 14 July. 


Case 2. On 3 June 1999, a nest at Cavendish Sands- 
pit (46.5028°N, 63.4322°W) was completed with four 
eggs. On 17 June, this nest was flooded and the eggs 
were found approximately | m from the nest in 10 cm 
of water. Both adults were present and appeared agi- 
tated. The real eggs were removed and clay eggs were 
placed in the nest. One severely cracked egg was dis- 
carded and the three undamaged eggs were returned 
to the nest once the water receded. One chick hatched 
on 30 June, and a second was observed on 1 July. The 
remaining egg was abandoned. Only one chick sur- 
vived to fledge and it was banded on 12 July. This 
chick was recaptured as a breeding male on the same 
beach in 2001. 


Case 3. On 17 June 1999, a nest with three eggs at 
Cavendish Sandspit (46.5047°N, 63.4483°W) was 
flooded without damage to the eggs. The eggs were 
found under 10 cm of water, more than | m from the 
nest. They were replaced with clay eggs. An adult was 
observed incubating the clay eggs approximately 
1.5 hours later. The real eggs were then replaced and 
both adults returned within 30 minutes. Only one 
adult was observed on subsequent visits and the nest 
was considered abandoned on 20 June. The eggs were 
collected on 22 June, candled, found to be viable and 
placed in an incubator. The eggs were candled again 
on 30 June, when further growth and movement was 
observed in all 3 eggs. The eggs were successfully 
fostered to another Piping Plover pair on 3 July, but 
were taken by a predator on 4 July. The eggs from 
this nest were not only flooded, but endured a 24-48 
hour lapse in incubation without any apparent decrease 
in viability. 

Case 4. A nest with three eggs was discovered on 
12 June 2001 at Sebim Beach in Sand Hills Prov- 
incial Park, Nova Scotia (43.5311°N, 65.5580°W),. 
The full clutch remained on 25 June, but the high tide 
had completely overwashed the nest and the sand 
remained wet. Only two eggs remained on 11 July, 
and incubation was confirmed on 13 July. One aban- 
doned egg remained in the nest on 16 July. On 18 


NOTES 


449 


July, a recently hatched chick (<Sdays of age) and 
one adult were observed. The development timeline 
indicates a protracted incubation period of at least 32 
days. Neither chick nor adults were observed after 18 
July, and the rearing period was too short for the chick 
to have fledged successfully. 

No information is available on the resistance of 
Piping Plover eggs to submersion (Haig 1992). Ward 
and Burger (1980) found that some embryos of Her- 
ring Gull (Larus argentatus) eggs could remain viable 
after immersion in cold seawater. As Piping Plover 
eggs have long been exposed to selection due to tidal 
flooding, they may be relatively resistant to the effects 
of inundation (Ward and Burger 1980). Piping Plovers 
exhibit strong parental tenacity to incubate and tend 
eggs (Prellwitz et al. 1995) and it is likely that their 
quick resumption of incubation of eggs after flooding 
increases the probability of successful hatching. 
These nest histories confirm that some Piping Plover 
embryos can survive immersion in seawater, and that 
replacing eggs washed out of recently flooded nests 
will increase the survival of individual clutches. 

Protracted incubation periods due to nest cooling 
have been recorded for Piping Plovers and other 
shorebirds (Cairns 1982). Therefore, eggs believed to 
have been immersed in seawater, but still present in 
the nest cup, should not be removed as long as the 
adults continue to incubate and the incubation period 
is shorter than 40 days. 


Acknowledgments 

We thank to Peter MacDonald, Amy Marsters, the 
field staff at Prince Edward Island National Park, 
Scott McBurney, Phil McCabe and Roland McCorm- 
ick. Canadian Wildlife Service, Parks Canada and the 
Nova Scotia Department of Natural Resources pro- 
vided funding and logistical support. 


Literature Cited 

Boyne, A., and D. Amirault. 1999. Habitat characteristics 
of Piping Plover nesting beaches in Nova Scotia, New 
Brunswick, and Prince Edward Island. Pages 84-85 in 
Proceedings, Piping Plovers and Least Terns of the Great 
Plains and nearby. Edited by K. F. Higgins, M. R. Brash- 
ier, and C. D. Kruse, South Dakota State University. Brook- 
ings, South Dakota, 132 pages. 

Burger, J. 1987. Physical and social determinants of nest 
site selection in Piping Plover in New Jersey. Condor 89: 
811-818. 

Cairns, W. E. 1982. Biology and behavior of breeding Pip- 
ing Plovers. Wilson Bulletin 94: 531-545. 

Cairns, W. E., and I. A. McLaren. 1980. Status of the Pip- 
ing Plover on the east coast of North America. American 
Birds 34: 206-208. 

Goossen, J. P., D. L. Amirault, J. Arndt, R. Bjorge, S. Boates, 
J. Brazil, S. Brechtel, R. Chiasson, G. N. Corbett, F. 
R. Curley, M. Elderkin, S. P. Flemming, W. Harris, L. 
Heyens, D. Hjertaas, M. Huot, B. Johnson, R. Jones, 
W. Koonz, P. Laporte, D. McAskill, R. I. G. Morrison, 
S. Richard, F. Shaffer, C. Stewart, L. Swanson, and E. 


450 


Wiltse. 2002. National Recovery Plan for the Piping 
Plover (Charadrius melodus). National Recovery Plan 
Number. 22. Recovery of Nationally Endangered Wildlife, 
Ottawa, 47 pages. 

Haig, S. M. 1992. The Piping Plover. Pages 1-18 in The birds 
of North America (2). Edited by A. Poole, P. Stettenheim, 
and F. Gill, American Ornithologists’ Union, Washington, 
Dic: 

Prellwitz, D. M., K. M. Erickson, and L. M. Osborne. 
1995. Translocation of Piping Plover nests to prevent nest 
flooding. Wildlife Society Bulletin 23: 103-106. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Sylvester, J. 1991. Privacy please! People and plovers flock 
to P.E.I.’s beaches, keeping them apart is a matter of the 
birds’ survival. Canadian Geographic 111(2): 37-40. 

United States Fish and Wildlife Service. 1996. Piping 
Plover (Charadrius melodus), Atlantic Coast Population, 
revised recovery plan. Hadley, Massachusetts. 258 pages. 

Ward, L. D., and J. Burger. 1980. Survival of herring gull 
and domestic chicken embryos after simulated flooding. 
Condor 82: 142-148. 


Received 29 May 2003 
Accepted 8 November 2004 


A Tribute to Loris Shano Russell, 1904-1998 


KEVIN L. SEYMOUR 


Department of Natural History, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario M5S 2C6 Canada; e-mail: 


kevins @rom.on.ca 


Seymour, Kevin L. 2004. A tribute to Loris Shano Russell. 1904-1998. Canadian Field-Naturalist 118(3): 451-464. 


It was somewhat daunting to try to write the story 
of a man as accomplished as Loris Shano Russell. On 
one hand, it is relatively easy to record or list the 
numerous publications and awards. On the other hand, 
it is much harder to see through all of the accom- 
plishments in order to glimpse something of the man 
himself. I got to know Loris only later in his life, as 
the one tasked with arranging his weekly visits to the 
Royal Ontario Museum (ROM), and so in this report 
I have relied on many who knew him earlier, in par- 
ticular, John E. Storer. 

Russell’s accomplishments were many. He was one 
of the last of the “old school” of palaeontologists, 
those who studied and published original findings in 
geology, stratigraphy, and both fossil invertebrates and 
vertebrates: a broad concept rather foreign to most of 
us in today’s specialized world. He was extremely 
organized, and was meticulous with everything he did, 
whether it was science, museology, ham and military 


radio operation, or administration. He was unfailingly 
polite and dignified, and always came to work wearing 
jacket and tie; about the worst name he ever called 
anyone in public was “stuffed shirt”, a phrase he 
reserved for H. F. Osborn. 

Loris died in July 1998, in his 95" year, and was 
predeceased by his beloved wife Grace, in March 1998. 
Their partnership of 60 years is the stuff of legends. 
They did everything together, from hosting museum 
dignitaries at their home, to attending antique markets, 
to working in the dirt and the heat in the badlands of 
Alberta collecting fossils (Figure 1). Grace never did 
learn very much about the fossils, but that didn’t matter 
to her at all — she was there for, and with, Loris. When 
he was hospitalised in 1990, Grace visited several times 
a week, as long as the Wheel-Trans vehicles were 
available to take her there, as she was then not very 
mobile herself. Their separation at this time must have 
been very difficult for both of them. Grace was’ an out- 


Ficure 1. Grace and Loris Russell, 10 June 1986, in the Red Deer River valley, at the site where the holotype of Edmon- 
tosaurus regalis was collected by Levi and C. H. Sternberg in 1912. Photo by Maurice Stefanuk. (Photo courtesy 


ROM Archives). 


45] 


452 


spoken fan of her husband, and she was as forthright 
with people as he was quiet and formal. Their differ- 
ences were marked, yet their partnership flourished. 
They had no children, and the topic was never dis- 
cussed with others. Many “Grace” stories exist; several 
are related in Churcher (2003). Francis Cook passed 
on the following story. At the NMC (National Museum 
of Canada, now the Canadian Museum of Nature) staff 
party for Loris when he was leaving and moving to the 
ROM, Grace invited everyone to visit them in their new 
quarters in Toronto and to use their swimming pool. 
Loris quietly yet respectfully deflected her enthusiasm 
by pointing out that they were moving to an apartment 
building and he was not sure that it included privileges 
for an infinite number of visitors to use the pool. 

Loris Shano Russell was born in April 1904, in 
Brooklyn, New York. His parents were Matilda Shano 
of Newfoundland, and Milan Winslow Russell of New 
York. Loris’ middle name came from his mother’s 
maiden name, of course, and Loris told me that “Shano” 
was an anglicised version of the French “Chenaud”. 
As for his unusual first name, Loris told me that his 
parents simply were looking for something different 
when they picked it; there was no family history to 
the name. 

In 1908, when Loris was four, his family moved to 
Calgary, Alberta, where he grew up. He attended both 
public and high school in Calgary, and must have had 
a keen early interest in science, judging by early photos 
of him (Figures 2 and 3). He attended the University 
of Alberta, Edmonton, and graduated with a B.Sc. in 
Geology in 1927; however, he had already started pub- 
lishing before graduation! He spent some time prospect- 
ing the Paskapoo Formation in Alberta near his fami- 
ly’s home: his first two reviewed papers (published in 
1926) are both on fossils of the Paskapoo. By this time 
he must have already encountered the Sternberg family 
of dinosaur collecting fame, as there is a 1923 photo 
of Loris excavating in the Red Deer River Valley, north 
of the Bleriot Ferry (Figure 4). Even at this early point 
in his career, his published papers were representative 
of his broad-ranging interests: one is on fossil mol- 
luscs, and the other on the fossil mammal Catopsalis. 
Fossils were not his only early interest, however. Before 
his first refereed papers on fossils, he had published a 
note on Alberta’s birds in 1923. 

At Princeton University, he studied under William 
Berryman Scott, the famous geologist and palaeon- 
tologist. W. A. Parks, then Head of the Department of 
Geology at the University of Toronto (UT), had 
wanted him to study at the UT, but Loris thought that 
it would be better to study with Scott, one of the very 
few European-trained professional vertebrate palaeon- 
tologists then teaching in North America. Loris was 
awarded his M.A. in 1929, and his Ph.D. in 1930, for 
a dissertation entitled “Stratigraphy and Paleontology 
of the Uppermost Cretaceous and Lower Tertiary 
Formations of Alberta”, a copy of which is in the ROM 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


FiGure 2. Loris Russell, 1920. Photographer unknown. 
(Photo courtesy ROM Archives). 


library. By graduation he had published at least a dozen 
papers, including papers on subjects as diverse as fossil 
pelecypods, gastropods, fish, turtles, dinosaurs, mar- 
supials and mammal tracks. These papers may have 
partly resulted from his summer work as a student 
assistant in 1925-1929 at the Research Council of 
Alberta in Edmonton. 

After graduating from Princeton in 1930 at the age 
of 26, he moved to Ottawa and served as Assistant 
Palaeontologist for the Geological Survey of Canada 
(GSC) until 1936, and an Assistant Geologist in 1937. 
During this period (1930-1937), he published over 
two dozen papers, again on a wide variety of topics: 
besides the requisite geological and stratigraphic works, 
there were a number of papers on fossil mammals 
and fresh-water molluscs, with smaller contributions 
on turtles, plesiosaurs and dinosaurs. Because of the 
utility of molluscs in biostratigraphy, many of his 
earlier works concentrated on these fossils. Of course 
his interests were not all palaeontological, and he 
joined the Ottawa Field-Naturalists’ Club in 1933. He 
later became an honorary member of this organization 
in 1972, after serving as Vice-President for 1954-1956, 
and as President for 1957-1958. 


2004 SEYMOUR: TRIBUTE TO LORIS SHANO RUSSELL 453 


Ficure 3. Loris Russell, 1922. Photographer unknown. 
(Photo courtesy ROM Archives). 


During his initial period in Ottawa (1930-1937) Loris 
met his future wife and constant companion of 60 years, 
Grace. Grace Evelyn LeFeuvre was eight years young- 
er than Loris. Her mother was born in Montreal of 
Irish stock, and her father had immigrated to Canada 
from Jersey, in the Channel Islands. Grace and Loris 
had met by arrangement of their mothers. While work- 
ing at the GSC in Ottawa, where he could take Grace 
out on dates, Loris was offered the position of Assistant 
Director of the Vertebrate Section of the then separate 
Royal Ontario Museum of Palaeontology (ROMP), 
in 1937. This position came about due to the death of 
W. A. Parks in 1936, and Parks’ protégé, Madeleine A. 
Fritz, simultaneously being appointed Assistant Direc- 
tor of the Invertebrate Section of ROMP. He consid- 
ered turning down the offer, because he did not want 
to be far from Grace. The only solution was marriage, 
and his proposal was to the point: “I’m not going there 
without you”. It appears he may have, at least briefly, 
because he moved to Toronto in 1937, and they were 
not married until 1938. Upon their marriage, Grace 
had to give up her nursing career, as only one income 
per household was allowed during the Depression. 

With the Assistant Directorship at the ROMP came 
an Assistant Professorship at the UT, in Palaeontology. 
However, this work was interrupted by the Second 
World War. With his ability as a ham radio operator 
(Figure 5), he served from 1942 to 1945 in the Royal 
Canadian Signal Corps. He first learned this skill in 
1922, an interest that he may have gotten from his father, 
who was a telegrapher for the railway (although his 
father had died earlier in 1911). At war’s end, he was 


Ficure 4. Loris Russell (right) and C. M. Sternberg (left) excavating an Edmontosaurus skeleton in the Red Deer River val- 
ley, 1923. Photo by J. E. Thurston. (Photo courtesy ROM Archives). 


454 


FicurE 5. Loris Russell, circa 1922. Photographer unknown. 
(Photo courtesy ROM Archives). 


transferred to the Reserve with the rank of Major. He 
continued an interest in communications, and collected 
early telegraph and other communications equipment, 
which since has been donated to the Museum of Sci- 
ence and Technology in Ottawa. In 1946 he was appoint- 
ed Director of the ROMP and in 1948 Associate Pro- 
fessor at the UT. During 1937-1950 he continued his 
studies of fossil vertebrates, producing some two dozen 
papers on fishes, dinosaurs, creodonts, titanotheres, hors- 
es, and mastodons, as well as others on the geology of 
Alberta, fossil gastropods, eurypterids and even living 
rattlesnakes! 

In 1950 came the offer to become Chief of the Zool- 
ogy Section at the NMC, an offer he could not resist. 
He and Grace returned to Ottawa, and remained there 
for the next 13 years. In 1956 he was appointed Direc- 
tor of the Natural History Branch of the NMC (Fig- 
ure 6), a post that he held until 1963. This time at the 
NMC was arguably the single most important part of 
his career as a research museum administrator, which 
he filled with steady competence, vision and perceptive 
guidance of staff activities. He hired several productive 
research and curatorial staff, including Wann Langston, 
Don McAllister and Arthur Clarke, who shared 
Loris’ professional interests in vertebrate palaeontol- 
ogy, ichthyology, and malacology, respectively. He 
profoundly influenced the direction of the Canadian 
natural sciences even outside his own fields of inter- 
est. For instance, in 1955 he suggested to invertebrate 
zoologist E. L. Bousfield that, as a staff member of a 
national institution, he might consider field studies 
on the Canadian Pacific coast and break from a previ- 
ous eight-year Ph.D. obsession with the Atlantic 
coast. This perceptive suggestion led to the discovery 
of a diverse, major fauna of amphipod crustaceans of 
which, during the next 30+ years, more than 200 
species and higher taxa were newly described. 

This period (1950-1963) must have been a very busy 
time for him, because in addition, in 1958 alone, he 
was appointed Acting Director of the Human History 
Branch at NMC, President of the Society of Verte- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


brate Paleontology, President of Section IV (Geolo- 
gy) of the Royal Society of Canada and he received 
an honorary LL.D. from the University of Alberta. 
He later became the President of the Canadian Muse- 
ums Association (CMA) in 1961, and was awarded 
the Willet G. Miller Medal from the Royal Society of 
Canada (RSC) in 1959. With respect to the RSC, at 
the time of his death in 1998 he was the most senior 
member (by seven years over three other elderly fel- 
lows) of the entire RSC roster of approximately 1500 
names, having been elected at the remarkable youth- 
ful age of 32 (in 1936), and with 62 years of mostly 
active participation in this select group of Canadian 
scientists. 

While at the NMC, Loris always had the deep respect 
of the staff, even though he had a bit of difficulty adjust- 
ing to the fact that they were no longer required to 
work Saturday mornings, as they had when he joined 
the GSC in the 1930s. When the NMC staff complained 
in winter about the cold and drafty Victoria Memorial 
Museum building where they all had offices, Loris qui- 
etly remarked that when Dr. Rand (Chief Zoologist 
from 1942-1947 and a very productive staff member) 
was there, he just put on an overcoat and went on work- 
ing. During 1950-1963, he wrote nearly 60 papers, with 
a broad range of topics: geology, eurypterids, molluscs, 
fishes, acanthodians, champsosaurs, carnivores, horses, 
and rabbits. Also during this time he produced more 
synthetic papers that included discussions of mam- 
malian migrations, continental zoology of the North 
American Pleistocene, the geological record of evolu- 
tion, as well as some on museology. These later papers 
were unlike anything he had written before, and spoke 
of a greater involvement in the museum community. 
Examples are several reports on television in museums, 
out-of-doors museums, plastics in the museum, and 
historical conservation along the St. Lawrence Seaway. 

Loris was intrigued by museums and was involved 
in the earliest days of the CMA. He was a forward- 
thinking museologist, whose central tenet could be 
summed up as: “Museums are doing their job when 
they are telling stories to the public”. This was not the 
prevailing thought in the 1960s, when the object was 
supposed to speak for itself, aided by lighting and gadg- 
ets, but it seems to be the popular notion once again, 
where story-centred galleries are becoming common. 

Although Loris may well have had a lifelong interest 
in material culture, it was during this period that this 
interest blossomed; being the President of the CMA 
from 1961-1963 and Acting Director of the Human 
History Branch of the NMC no doubt spurred it on. 
Constrained by a lack of time to do much palaeonto- 
logical research while handling administration, Loris 
decided he would try to apply scientific methods to 
some research in material history, as an experiment. 
He chose oil lamps as a subject for research, perhaps 
as a manifestation of industrial development coupled 
with social history, and it kept him “occupied and 


2004 


broke” for years. Loris and Grace visited antique shops 
together, often in small towns on their way to do palae- 
ontological fieldwork. Some of these adventures are 
described in his 1969 Rotunda article. His habit of 
meticulously documenting and labelling everything, 
plus his deep-seated love of a good story (he claimed 
it came from growing up in the “Wild West’), put 
him in the position to do ground-breaking research in 
material culture. Loris developed a superb collection 
of well-documented lamps, which were later kept in 
glass display cases in their Toronto apartment; these 
have since been donated to the ROM. Several books, 
A Heritage of Canadian Light (1968), Handy Things to 
Have Around the House (1979), and Every Day Life 
in Colonial Canada (1980) resulted from this research. 
These are still standard references today; indeed, A 
Heritage of Light was reprinted in February 2003 by 
the University of Toronto Press. He became a speaker 
in demand at various material culture conferences 
over the next few decades. 

In 1963, Loris left NMC under some controversy. 
Upheaval in the administrative ranks of the NMC made 
for some messy politics, which Loris did not care for. 
Returning to Toronto and the ROM, he became the 
Head of the Life Sciences Division at the ROM, and 
a year later, he filled the newly created position of 
Chief Biologist. By that time, the five former Royal 
Ontario Museums (zoology, palaeontology, mineral- 
ogy, geology and archaeology) had been amalgamat- 
ed into a single institution under one Director. Much 
of his continued success at ROM was due to Anne 
Liebeck, Loris’ able secretary (Figure 7). She guarded 
the entrance to Loris’ office and answered his telephone, 
so that he could continue his studies virtually without 
interruption. In 1964, he received a Diploma with 
Distinction from the Museums Association of Great 
Britain. With Loris’ return to Toronto came a profes- 
sorship in the Department of Geology at the UT. He 
took on three Ph.D. students before retiring: John Stor- 
er (graduated in 1970), and Paul Ramaekers and Mark 
Wilson, both of whom graduated in 1974. Loris offi- 
cially retired in 1970, when he was appointed Professor 
Emeritus at the UT in 1970, and Honorary Curator at 
the ROM in 1971. Loris continued to serve the scien- 
tific community after retirement — he was elected in 
1971 to the Presidency of the Royal Canadian Insti- 
tute, and in 1972 to the Presidency of the Internation- 
al Palaeontological Union, a four-year term. He pub- 
lished another 36 papers during this time, with a 
familiar breadth of topics. Several papers hinted of 
things to come: his museology papers concentrated 
on lighting and lamps, and his articles on “Tertiary 
Mammals of Saskatchewan, Part 1” and “The Great 
Saskatchewan Mouse Mine” began a series of papers 
on these important faunas from Saskatchewan. As 
well, Loris’ paper on “Body temperature of dinosaurs 
and its relationship to their extinction” in 1965 marks 
the first, and often overlooked, discussion in the Eng- 


SEYMOUR: TRIBUTE TO LORIS SHANO RUSSELL 


FiGure 6. Loris Russell, 8 November 1956, as Head of the 
Department of Natural History, National Museum of 
Canada. Photographer unknown. (Photo courtesy of 
the CMN Archives, negative #J4171). 


lish scientific literature of what was later to become a 
revolution in thinking: dinosaurs as active “warm- 
blooded” animals. Grace often cited this paper as one 
of the reasons she was so proud of “her hubby”. It 
was not a secret wish of Loris’ that perhaps Bob 
Bakker and John Ostrom (the oft-cited originators of 
this hypothesis) and others might have given him 
more credit for this early insightful work. 

His official retirement only meant less administra- 
tion. He continued fieldwork in Alberta each summer 
with Grace and others, well into his eighties, until at 
least 1988, supported by NSERC (Natural Science 
and Engineering Research Council of Canada) grants. 
These grants and the subsequent fieldwork meant that 
he actively published papers until about 1990, with 
over 40 contributions since retirement. He continued 
to come into his ROM office daily from 1971 until 
about 1990, when he first entered hospital for a hip- 
replacement operation. He never left hospitals after 
the first operation. From 1994 until 1997, while still 
wheelchair-bound due to a second failed hip replace- 
ment operation, I arranged that he visit ROM one day 
a week (except during winters) using Wheel-Trans 
services. During these visits he worked on his last 
manuscript, concerning the biostratigraphy of the 
Horseshoe Canyon Formation. Although this paper 
was never published, staff at the Royal Tyrrell Museum 
of Palaeontology and the Canadian Museum of Nature 
plan to publish a paper in the near future on the bios- 
tratigraphy of the Horseshoe Canyon Formation rec- 


456 


ognizing Loris’ essential contributions by including him 
as a posthumous co-author. Many important palaeon- 
tological contributions came from this post-retirement 
period, for example, the series of papers on the Ter- 
tiary Mammals of Saskatchewan (Parts 2 through 7). 
More awards came late in his life, in particular, the 
Canadian Silver Jubilee Medal in 1978, the Billings 
Medal from the Geological Association of Canada in 
1984 and the Romer-Simpson Medal from the Society 
of Vertebrate Paleontology in 1992. 

As Tokaryk (1998) noted, much of Loris’ scientific 
work was accomplished solo, judging by the rarity of 
co-authored publications (only 10 out of more than 
200 papers published over 70 years have co-authors). 
This was partly due to the paucity of other Canadian 
palaeontologists at the time (although he was ably 
assisted, or worked with, a number of others in the field), 
but mostly to do with the fact that he could handle 
both the geological and palaeontological parts of his 
chosen projects. A measure of the impact of his work 
can be taken by counting the number of his papers that 
were abstracted in the German abstract series Palaeon- 
tologishes Zentralblatt (after 1950 called Zentralblatt 
fiir Geologie und Palaontologie, Teil 2) or Neues Jahr- 
buch fiir Mineralogie, Geologie und Paliontologie 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


4 


Ficure 7. Loris Russell and Anne Liebeck at ROM, 1971. Photo by L. R. Warren. (Photo courtesy ROM Photography). 


(after 1942 called Zentralblatt fiir Mineralogie, Geolo- 
gie und Palaontologie). Between 1927 and 1989, I 
found 68 Russell papers had been abstracted, by a 
total of 15 different reviewers (besides those who wrote 
anonymously), although more than half of this total 
were written by Jaworski or Wenz on molluscs and 
von Huene, primarily on dinosaurs. 

He was an adventuresome scientist, always willing 
to strike off in a new direction. While on holiday with 
Grace in Hawaii, he collected some fossil land snails 
at the famous Diamond Head locality of picture post- 
card fame. A drawer of this material remains at the 
ROM, labelled and researched, although he never did 
publish on them. Nor was he afraid of being wrong, 
or did he mind greatly if new research invalidated a 
few of his ideas: that was the way science worked. As 
long as the work was careful and the facts were straight, 
then this was fine with him, an admirable quality in a 
scientist! 

Another quality that many people noticed in Loris 
was his phenomenal memory for field localities. He 
really could find fossil localities he hadn’t seen in 40 
years. His successes were legion, so the few exceptions 
stood out, and yet all of these exceptions seemed to 
have complicating factors. For instance, he was frus- 


2004 


trated about not being able to relocate the Calgary 
Paskapoo Formation localities he collected as a young 
student, only to realise that housing developments 
had probably covered the area. He tried many times 
to find Brown’s (1914) Erickson’s Landing locality. 
Although in Russell (1929) he reported relocating this 
locality, Krause (1978) expressed some doubt about 
this, and Loris must have harboured some doubts 
himself. Eventually, as reported by Fox (1990), Loris 
concluded that there was no way to determine from 
exactly which level the slump block containing the fos- 
sils was derived, and so he never was able to collect 
significant additional material. 

Certainly Loris had a happy outlook on life, and he 
taught by example, both in the field and in the labora- 
tory. He was an inspiring influence to all who worked 
with him. He was also the kind of person who would 
retreat to his workshop to solve design problems, per- 
haps a legacy of his ham radio days. He designed a 
unique machine for feeding a thin stream of washed 
fossil concentrate onto a rubber belt (Figure 8), the 
movement of which was controlled by a foot pedal. 
The belt passed under a microscope, allowing him to 
focus on, and select out, any fossils of interest with 
his free hands. Uninteresting concentrate rotated off 
the belt and into a box of scrap. We have preserved this 
machine at the ROM; perhaps someday others will 
use it. It was known by words beginning with the let- 
ter ‘M’, such as ‘Miraculous Moving Miocene Mouse 
Machine’. For work on site at the Kleinfelder Farm 
locality in Saskatchewan (appropriately enough called 
the ‘Mouse Mine’), he also designed two rotary sieves, 
described in his 1970 Rotunda article. 

Although Dr. Russell (as he was known) seemed a 
little stiff and intimidating with some people, he actu- 
ally had a wry yet somewhat playful sense of humour 
that surfaced quietly with those he knew well. He was 
quite amused by the ironies of growing older. On a 
couple of occasions he observed: “We used to call 
Scott and Osborn and their generation the ‘old boys’, 
and look how things have turned out now”. He would 
tell the story of being a young geological assistant in 
the 1920s when one evening he got some sort of 
buzzing insect stuck in his ear. On asking another 
assistant, who was a medical student, what he should 
do, he was told “Well, take it out of there!”. Loris’ 
father was a religious fundamentalist, so Loris learned 
the Bible while young. In his later years he was not 
religious, but appreciated and was amused by the fun- 
damentalist side of Western life. He liked to recount 
Charlie Sternberg’s remark “My, aren’t we clever’, 
when William “Bible Bill” Aberhart (Dean of the 
Calgary Prophetic Bible Institute) stated in a radio 
sermon that palaeontologists were actually manufac- 
turing fake dinosaur bones in seclusion in the bad- 
lands. At a gathering for an NMC staff member about 
to be married, Francis Cook noticed Loris was drink- 
ing milk, whereas the rest of those in attendance were 


SEYMOUR: TRIBUTE TO LORIS SHANO RUSSELL 


457 


FiGure 8. Loris Russell with the Marvellous Moving Miocene 
Mouse Machine, 1981. Photographer unknown. (Photo 
courtesy ROM Photography). : 


not. The host, when this was mentioned, assured Fran- 
cis that Loris was drinking milk only because he had 
an ulcer, but that he had laced it with whiskey, there- 
by obeying doctor’s orders yet joining his colleagues. 
He always found his own way. 

Several obituaries already have been written (Har- 
ington 1998; Shaul 1998; Sues 1998; Tokaryk 1998; 
Churcher 2003), and two appreciations (Swinton 1976; 
Churcher 1993), with many additional biographical 
details. Swinton (1976) also included a fairly complete 
Russell bibliography up to 1976. A more complete 
bibliography is included herein. 

The Russell papers, including diaries, field notes, 
photos, films (several different kinds), correspondence 
and other records, were inventoried by Boden (1999) 
and Baltovich (2001), and are placed in the ROM 
archives. His slides and reprints of scientific articles 
are stored in the Section of Palaeobiology, Depart- 
ment of Natural History, ROM. Hopefully, someone 
in the future will take advantage of this material (par- 
ticularly the diaries and films) and write a book on 
Loris Russell. 


Acknowledgments 

For personal information: John Storer (Yukon 
Department of Tourism and Culture, Whitehorse), 
Rufus Churcher (retired from the UT), Tony Flynn 
(Seneca College, Toronto) and Grace’s niece Barbara 


458 


Rager, of Ottawa. Francis Cook, E. L. Bousfield, Dick 
Harington and Richard Day, past and present NMC 
(now CMN) staff, read an earlier draft and contributed 
many useful comments and additions. In particular, 
Richard Day located many bibliographic citations for 
abstracts (and this started me looking for more!). For 
salvaging the Russell diaries for the ROM: Tony Flynn. 
For access to the ROM Archives, and for their interest 
in the Russell papers: Julia Matthews and Sharon Hick. 
For bibliographic assistance: Arthur Smith. For pho- 
tographic assistance: Alison Murray and Peter Fenton. 
I would like to most sincerely thank all of these people 
for their assistance with this tribute; without them it 
simply would not have been completed. 


Documents and Literature Cited 

Baltovich, R. 2001. Report on inventory project [adden- 
dum]: the Loris Shano Russell papers. Unpublished 
report, 18 pages. [on file in ROM Library and ROM Sec- 
tion of Palaeobiology, Department of Natural History]. 

Boden, S. 1999. Report on inventory project: the Loris 
Shano Russell papers. Unpublished report, 31 pages. [on 
file in ROM Library and ROM Section of Palaeobiology, 
Department of Natural History]. 

Brown, B. 1914. Cretaceous-Eocene correlation in New 
Mexico, Wyoming, Montana, Alberta. Geological Soci- 
ety of America Bulletin 25: 355-380. 


Loris Shano Russell Bibliography 
Compiled by KEVIN L. SEYMOUR 


Included are books, book reviews, scientific arti- 
cles and those on material culture, but probably not 
all abstracts. Abstracts or reviews of Russell’s work 
that were written by others are not included, but many 
are on file in the Section of Palaeobiology, Depart- 
ment of Natural History, ROM. Within any year, the 
articles are not necessarily listed in strict chronologi- 
cal order. For those papers with co-authors, all authors 
are listed; if no author is listed, the paper was authored 
solely by L. S. Russell. Copies of all publications are 
on file in the Section of Palaeobiology, Department 
of Natural History, ROM. 


1923 Alberta and its birds. Bird-Lore 25(5): 348-350. 

1926a Technique. American Journal of Science 11: 498-500. 

1926b A new species of the genus Catopsalis Cope from the 
Paskapoo Formation of Alberta. American Journal 
of Science 12: 230-234. 

1926c Mollusca of the Paskapoo Formation in Alberta. Trans- 
actions of the Royal Society of Canada, Third Series, 
20 (Section 4): 207-220, Plates 1-3. 

1928a Didelphiidae from the Lance beds of Wyoming. Jour- 
nal of Mammalogy 9: 229-232. 

1928b A new fossil fish from the Paskapoo beds of Alberta. 
American Journal of Science 15: 103-107. 

1928c R. L. Rutherford and L. S. Russell. Mammal tracks 
from the Paskapoo beds of Alberta. American Jour- 
nal of Science 15: 262-264. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Churcher, C. S. 1993. Loris Shano Russell. (Eulogy for) 
Romer Simpson Prize. Society of Vertebrate Paleontol- 
ogy News Bulletin 157: 26-27. 

Churcher, C. S. 2003. Loris Shano Russell, 1904-1998. 
Proceedings of the Royal Society of Canada for 2001, 
Sixth series, 12: 265-270. 

Fox, R. C. 1990. The succession of Paleocene mammals in 
western Canada. Pages 51-70 in Dawn of the Age of 
Mammals in the northern part of the Rocky Mountain 
Interior, North America. Edited by T. M. Bown and K. D. 
Rose. Geological Society of America Special Paper 243. 

Harington, C. R. 1998. In memoriam: Loris Shano Russell 
1904-1998. Trail and Landscape 32(4): 166-169. 

Krause, D. W. 1978. Paleocene primates from western Can- 
ada. Canadian Journal of Earth Sciences 15: 1250-1271. 

Russell, L. S. 1929. Paleocene vertebrates from Alberta. 
American Journal of Science 17: 162-178. 

Shaul, S. 1998. Shining a light on the dim past: lives lived - 
Loris Shano Russell. Globe and Mail, August 12: A18. 
Sues, H.-D. 1998. Loris Shano Russell, 1904-1998. Society 
of Vertebrate Paleontology News Bulletin 174: 81-83. 
Swinton, W. E. 1976. Loris Shano Russell: an appreciation. 
Pages 3-13 in Athlon: essays on palaeontology in honour 
of Loris Shano Russell. Edited by C. S. Churcher. Royal 
Ontario Museum, Life Sciences Miscellaneous Publica- 

tion. 

Tokaryk, T. T. 1998. One of the last of the early giants. 
Geology Today 14(6): 226-228. 


1928d The genera Kindleia and Stylomyleodon. American 
Journal of Science 15: 264. 

1929a Paleocene vertebrates from Alberta. American Journal 
of Science 17: 162-178. 

1929b The validity of the genus Stylomyleodon. American 

Journal of Science 17: 369-371. 

Upper Cretaceous and Lower Tertiary Gastropoda from 

Alberta. Transactions of the Royal Society of Cana- 

da, Third Series, 23 (Section 4): 81-90, Plate 1. 

1930a Early Tertiary mammal tracks from Alberta. Transac- 
tions of the Royal Canadian Institute 17: 217-221, 
Plates 7-11. 

1930b A new species of Aspideretes from the Paskapoo For- 
mation of Alberta. American Journal of Science 20: 
27-32. 

1930c Upper Cretaceous dinosaur faunas of North America. 
Proceedings of the American Philosophical Society 
69: 133-159. 

1930d Fresh-water plesiosaurs. Bulletin of the Geological 
Society of America 41: 198. [Abstract]. 

1930e Stratigraphy and paleontology of the Uppermost Cre- 
taceous and Lower Tertiary Formations of Alberta. 
Ph.D. thesis, Princeton University, 91 pages. 

1931la Mollusca from the Upper Cretaceous and Lower Ter- 
tiary of Alberta. Transactions of the Royal Society of 
Canada, Third Series, 25 (Section 4): 9-19, Plates 1-2. 

1931b Fresh-water plesiosaurs. Pan-American Geologist 53: 
153. [Abstract]. 

193lc Fresh-water plesiosaurs. Canadian Field-Naturalist 
45: 135-137. 


1929c 


2004 


1931d 
193 le 


1932a 


1932b 


1932c 


1932d 


1932e 


1932f 


1932g 


1933a 


1933b 


1933c 
1933d 


1933e 


1933f 


1933¢ 


1934a 


1934b 


1934c 


1934d 


1934e 


1934f 


1934¢ 


SEYMOUR: TRIBUTE TO LORIS SHANO RUSSELL 


Early Tertiary Mollusca from Wyoming. Bulletins of 
American Paleontology 18(64): 1-38, Plates 1-4. 
Devonian fishes of eastern Canada. Bulletin of the 
Geological Society of America 42:361. [Abstract] 
Mollusca from the McMurray Formation of Northern 
Alberta. Transactions of the Royal Society of Canada, 
Third Series, 26 (Section 4): 1-7, Plate 1. 

The Cretaceous-Tertiary transition of Alberta. Trans- 
actions of the Royal Society of Canada, Third Series, 
26 (Section 4): 121-156. 

New data on the Paleocene mammals of Alberta, Cana- 
da. Journal of Mammalogy 13: 48-54. 

On the occurrence and relationships of the dinosaur 
Troé6don. Annals and Magazine of Natural History, 
Series 10, 9: 334-337. 

New species of Mollusca from the St. Mary River For- 
mation of Alberta. Canadian Field-Naturalist 46: 80-81. 
Stratigraphy and structure of the eastern portion of the 
Blood Indian reserve, Alberta. Geological Survey of 
Canada, Summary Report 1931B: 26-38. 

Fossil non-marine Mollusca from Saskatchewan. Trans- 
actions of the Royal Canadian Institute 18: 337-341, 
Plate 1. 

A new species of Merychippus from the Miocene of 
Saskatchewan. Canadian Field-Naturalist 47: 11. 

L. S. Russell and R. T. D. Wickenden. Discovery of 
an Upper Eocene mammalian fauna in Southern 
Saskatchewan. Bulletin of the Geological Society of 
America 44: 199. [Abstract]. 

Peking Man. The Civil Service Review 5: 104. 

L. S. Russeil and R. T. D. Wickenden. An Upper 
Eocene vertebrate fauna from Saskatchewan. Trans- 
actions of the Royal Society of Canada, Third Series, 
27 (Section 4): 53-66, Plate 1. 

L. S. Russell and R. T. D. Wickenden. Upper Eocene 
vertebrate fauna from Saskatchewan, Canada. Proceed- 
ings of the Royal Society of Canada, Third Series, 
27 (Appendix B): 143. [Abstract by title only]. 
Revision of the Lower Oligocene vertebrate fauna of 
the Cypress Hills, Saskatchewan. Proceedings of the 
Royal Society of Canada, Third Series, 27 (Appen- 
dix B): 144. [Abstract by title only]. 

Early Tertiary mammal tracks from Alberta. Palaeon- 
tologisches Zentralblatt 2: 321. [Abstract]. 
Reclassification of the fossil Unionidae (fresh-water 
mussels) of Western Canada. Canadian Field-Naturalist 
48: 1-4. 

Pleistocene and post-Pleistocene molluscan faunas of 
Southern Saskatchewan. Canadian Field-Naturalist 
48: 34-37. 

New fossil fresh-water Mollusca from the Cretaceous 
and Paleocene of Montana. Journal of the Washing- 
ton Academy of Sciences 24: 128-131. 

Discovery of Middle Eocene Mammalia in British 
Columbia. Proceedings of the Geological Society of 
America for 1933: 368. [Abstract]. 

Restoration of the horned dinosaur Chasmosaurus. 
Proceedings of the Geological Society of America for 
1933: 368. [Abstract]. 

Revision of the Lower Oligocene vertebrate fauna of 
the Cypress Hills, Saskatchewan. Transactions of the 
Royal Canadian Institute 20: 49-67, Plates 7-10. 
Fossil turtles from Saskatchewan and Alberta. Pro- 
ceedings of the Royal Society of Canada, Third Series, 
28 (Appendix C): 113. [Abstract]. 


1934h 


19341 


1936b 


1936c 


1936d 


1937a 


1937b 


1937c 


1937d 


1937e 


1937f 


1938a 


1938b 


459 


Fossil turtles from Saskatchewan and Alberta. Trans- 
actions of the Royal Society of Canada, Third Series, 
28 (Section 4): 101-110, Plates 1-6. 

An Upper Eocene vertebrate fauna from Saskatchewan. 
Palaeontologisches Zentralblatt 5: 43. [Abstract]. 

A Middle Eocene mammal from British Columbia. 
American Journal of Science 29: 54-55. 
Musculature and function in the Ceratopsia. Bulletin 
of the National Museum of Canada 77: 39-48. 
Fauna of the Upper Milk River beds, Southern Alberta. 
Proceedings of the Royal Society of Canada, Third 
Series, 29 (Appendix B): 99. [Abstract]. 

Fauna of the Upper Milk River beds, Southern Alberta. 
Transactions of the Royal Society of Canada, Third 
Series, 29 (Section 4): 115-128, Plates 1-5. 

FE. Jj. Fraser, F. H. McLearn, L. S. Russell, P. S. War- 
ren, and R. T. D. Wickenden. Geology of Southern 
Saskatchewan. Geological Survey of Canada Mem- 
oir 176: 1-137, Plates 1-5. 

A Middle Eocene mammal from British Columbia. 
Palaeontologisches Zentralblatt 6: 378. [Abstract]. 
Plesiosaur from the Upper Cretaceous of Manitoba. 
Proceedings of the Geological Society of America 
for 1934: 378. [Abstract]. 

Dinosaur restoration group in the National Museum 
of Canada. Proceedings of the Geological Society of 
America for 1934: 378. [Abstract]. 

A plesiosaur from the Upper Cretaceous of Manitoba. 
Journal of Paleontology 9: 385-389, Plates 44-46. 
Revision of the Lower Oligocene vertebrate fauna of 
the Cypress Hills, Saskatchewan. Palaeontologisches 
Zentralblatt 7: 49. [Abstract]. 

Oil and gas possibilities along Milk River, south- 
eastern Alberta. Geological Survey of Canada Paper 
36-12: 1-24. 

Second multituberculate from the Belly River Forma- 
tion of Alberta. Proceedings of the Geological Society 
of America for 1935: 403. [Abstract]. 

New and interesting mammalian fossils from western 
Canada. Transactions of the Royal Society of Canada, 
Third Series, 30 (Section 4): 75-80, Plate 1. 

A plesiosaur from the Upper Cretaceous of Manitoba. 
Palaeontologisches Zentralblatt 8: 327. [Abstract]. 
Revision of the geology of the southern Alberta plains. 
Transactions of the Canadian Institute of Mining and 
Metallurgy 40: 185-196. 

Preliminary Report. Del Bonita area, southern Alber- 
ta. Geological Survey of Canada, Paper 37-10: 1-12. 
L. S. Russell and J. C. Sproule. Preliminary Report. 
Geology of the vicinity of Taber, Alberta. Geological 
Survey of Canada, Paper 37-14: 1-7. 

New non-marine Mollusca from the Upper Cretaceous 
of Alberta. Proceedings of the Royal Society of Cana- 
da, Third Series, 31 (Appendix B): 143. [Abstract]. 
New non-marine Mollusca from the Upper Cretaceous 
of Alberta. Transactions of the Royal Society of Cana- 
da, Third Series, 31 (Section 4): 61-67, Plate 1. 
Fossil turtles from Saskatchewan and Alberta. Palaeon- 
tologisches Zentralblatt 9: 373. [Abstract]. 
Rattlesnakes in Alberta. Canadian Geographical Jour- 
nal 16(1): 33-41. 

New species of Gastropoda from the Oligocene of 
Colorado. Journal of Paleontology 12: 505-507. 


460 


1938c 


1938d 


1938e 


1938f 


1938¢ 


1938h 


1939a 


1939b 


1939¢ 


1939d 


1939e 


1940a 


1940b 


1940c 


1940d 


1940e 


1940f 


1940¢ 


1940h 


19401 


THE CANADIAN FIELD-NATURALIST 


Complete skull of a titanothere from the Lower Oligo- 
cene of Saskatchewan. Bulletin of the Geological Soci- 
ety of America 49: 1920. [Abstract]. 

The skull of Hemipsalodon grandis, a giant Oligocene 
creodont. Proceedings of the Royal Society of Canada, 
Third Series, 32 (Appendix D): 146. [Abstract]. 
Origin of the sandstone dykes of South-eastern Alberta. 
Proceedings of the Royal Society of Canada, Third 
Series, 32 (Appendix D): 146. [Abstract]. 

The skull of Hemipsalodon grandis, a giant Oligocene 
creodont. Transactions of the Royal Society of Canada, 
Third Series, 32 (Section 4): 61-66, Plates 1-5. 

New species of Gastropoda from the Oligocene of 
Colorado. Proceedings of the Geological Society of 
America for 1937: 288. [Abstract]. 

Skull of Hemipsalodon grandis Cope. Proceedings 
of the Geological Society of America for 1937: 288. 
[Abstract]. 

Land and sea movements in the Late Cretaceous of 
Western Canada. Proceedings of the Royal Society of 
Canada, Third Series, 33 (Appendix C): 198. [Abstract]. 
Edmontonia rugosidens (Gilmore), an armoured 
dinosaur from the Belly River series of Alberta. Pro- 
ceedings of the Royal Society of Canada, Third 
Series, 33 (Appendix C): 200. [Abstract]. 

Land and sea movements in the Late Cretaceous of 
Western Canada. Transactions of the Royal Society 
of Canada, Third Series, 33 (Section 4): 81-99. 
Sclerotic ring in the skull of Lambeosaurus. Bulletin 
of the Geological Society of America 50: 1966. 
[Abstract]. 

Notes on the occurrence of fossil fishes in the Upper 
Devonian of Maguasha, Quebec. Contributions of the 
Royal Ontario Museum of Palaeontology 2: 1-10, 
Plate 1. 

The sclerotic ring in the Hadrosauridae. Contributions 
of the Royal Ontario Museum of Palaeontology 3: 
1-7, Plates 1-2. 

Studies of the Tertiary gravel deposits of Southern 
Saskatchewan. Proceedings of the Royal Society of 
Canada, Third Series, 34 (Appendix D): 158. [Abstract]. 
Titanotheres from the Lower Oligocene Cypress Hills 
Formation of Saskatchewan. Proceedings of the Royal 
Society of Canada, Third Series, 34 (Appendix D): 
159. [Abstract]. 

Titanotheres from the Lower Oligocene Cypress 
Hills Formation of Saskatchewan. Transactions of 
the Royal Society of Canada, Third Series, 34 (Sec- 
tion 4): 89-100, Plates 1-5. 

Edmontonia rugosidens (Gilmore), an armoured 
dinosaur from the Belly River series of Alberta. Uni- 
versity of Toronto Studies, Geological Series 43: 1- 
28, Plates I to VIII. 

Geology of the Southern Alberta plains. Part 1, 
Stratigraphy and structure. Geological Survey of 
Canada Memoir 221: 1-128, 3 maps. 

Micrichnus tracks from the Paskapoo Formation of 
Alberta. Transactions of the Royal Canadian Insti- 
tute 23(49): 67-74, Plates 1-2. 

Discovery of a marine fauna in Eastend Formation 
of Saskatchewan, Canada. Bulletin of the Geologi- 
cal Society of America 51: 1976. 

New genus of Gastropoda, probable ancestor of the 
Grangerellidae from the Upper Cretaceous of Alberta. 


1941a 


1941b 


1942a 


1942b 


1943a 


1943b 


1946a 


1946b 


1946c 


1946d 


1946e 


1947a 


1947b 


1947c 


1948a 


1948b 


1948c 


1948d 


1949a 


1949b 


Vol. 118 


Bulletin of the Geological Society of America 51: 
1976. 

Prograngerella, a new ancestral land snail from the 
Upper Cretaceous of Alberta. Journal of Paleontol- 
ogy 15: 309-311. 

Pleistocene horse remains from Saskatchewan. Pro- 
ceedings of the Royal Society of Canada, Third Series, 
35 (Appendix C): 188. [Abstract]. 

The structure of the crest in the dinosaur Parasaurolo- 
phus. Proceedings of the Royal Society of Canada, 
Third Series, 36 (Appendix C): 146. [Abstract]. 

The application of geology to strategy and tactics. 
Proceedings of the Royal Society of Canada, Third 
Series, 36 (Appendix C): 146. [Abstract]. 
Pleistocene horse teeth from Saskatchewan. Journal 
of Paleontology 17: 110-114. 

Marine fauna of the Eastend Formation of Saskat- 
chewan. Journal of Paleontology 17: 281-288, Plates 
47-49. 

The crest of the dinosaur Parasaurolophus. Contri- 
butions of the Royal Ontario Museum of Palaeontol- 
ogy 11: 1-5. 

Geophysical aspects of land-mine detection. Proceed- 
ings of the Royal Society of Canada, Third Series, 
40 (Appendix C): 170. [Abstract]. 

The lower jaw of the theropod dinosaur Troédon. 
Proceedings of the Royal Society of Canada, Third 
Series, 40 (Appendix C): 171. [Abstract]. 
Preliminary report on the stratigraphy of the Gaspé 
limestone series, Forillon Peninsula, Cap des Rosiers 
township, County of Gaspé South. Province of Quebec 
Department of Mines, Bureau of Geological Surveys, 
Preliminary Report 195: 1-14. 

Rapport préliminaire sur la stratigraphie des séries de 
calcaires de Gaspé, péninsule de Forillon, canton de 
Cap des Rosiers, Comté de Gaspé Sud. Province de 
Québec, Ministére des Mines, Service de la Carte 
Géologique, Rapport préliminaire 195: 1-16 [French 
translation of above]. 

Late Cretaceous and Early Tertiary correlation in 
Alberta and Saskatchewan. Bulletin of the Geologi- 
cal Society of America 58: 1223. [Abstract]. 
Post-glacial mastodon remains from southwestern 
Ontario. Bulletin of the Geological Society of Amer- 
ica 58: 1223. [Abstract]. 

A new locality for fossil fishes and eurypterids in the 
Middle Devonian of Gaspé, Quebec. Contributions 
of the Royal Ontario Museum of Palaeontology 12: 
1-6. 

A Middle Paleocene mammal tooth from the foothills 
of Alberta. American Journal of Science 246: 152- 
156, Plate 1. 

The dentary of Troddon, a genus of theropod dino- 
saurs. Journal of Paleontology 22: 625-629. 
Post-glacial occurrence of mastodon remains in south- 
western Ontario. Transactions of the Royal Canadian 
Institute 27(57): 57-64. 

Faunal facies in the Late Cretaceous marine deposits 
of western Canada. Bulletin of the Geological Society 
of America 59: 1348-1349. [Abstract]. 

The relationships of the Alberta Cretaceous dinosaur 
“Laosaurus” minimus Gilmore. Journal of Paleon- 
tology 23: 518-520. 

Preliminary Report. The geology of the southern 
part of the Cypress Hills, southwestern Saskatchewan. 


2004 


1950a 


1950b 


1950c 


195la 


1951b 


1951c 


1951d 


195le 


1951f 


1952a 


1952b 
1952c 
1952d 


1952e 


1952f 


1952¢ 


1952h 


1953a 


1953b 


1953¢ 


1953d 


1953e 


SEYMOUR: TRIBUTE TO LORIS SHANO RUSSELL 


Saskatchewan Geological Survey, Petroleum Geolo- 
gy Series 1: 1-56. 

Correlation of the Cretaceous-Tertiary transition in 
Saskatchewan and Alberta. Bulletin of the Geologi- 
cal Society of America 61: 27-42. 

The Tertiary gravels of Saskatchewan. Proceedings 
of the Royal Society of Canada, Third Series, 44 
(Appendix F): 226. [Abstract]. 

The Tertiary gravels of Saskatchewan. Transactions 
of the Royal Society of Canada, Third Series, 44 
(Section 4): 51-59. 

Acanthodians of the Upper Devonian Escuminac For- 
mation, Maguasha, Quebec. Annals and Magazine 
of Natural History, Series 12, 4: 401-407. 
Personalities in paleontology (Charles M. Sternberg). 
News Bulletin of the Society of Vertebrate Paleon- 
tology 31: 28-30. 

Bobasatrania? canadensis (Lambe), a giant chon- 
drostean fish from the Rocky Mountains. Annual 
Report of the National Museum of Canada for the fis- 
cal year 1949-50, Bulletin 123: 218-224, Plates 45-46. 
Age of the Front-Range deformation in the North 
American Cordillera. Proceedings of the Royal 
Society of Canada, Third Series, 45 (Appendix D): 
201. [Abstract]. 

Age of the Front-Range deformation in the North 
American Cordillera. Transactions of the Royal Soci- 
ety of Canada, Third Series, 45 (Section 4): 47-69. 
Land snails of the Cypress Hills and their signifi- 
cance. Canadian Field-Naturalist 65: 174-175. 

The Northeast Conference of the American Associa- 
tion of Museums, Cooperstown, New York, 19th & 
20" October, 1951. Bulletin of the Canadian Muse- 
ums Association 5(1): 7-12. 

Succession duties and museum bequests. Bulletin of 
the Canadian Museums Association 5(1): 13-14. 
Television and museums — an interim report. Bulletin 
of the Canadian Museums Association 5(1): 15-16. 
Out-of-doors museums. Bulletin of the Canadian 
Museums Association 5(2): 9-12. 

A new species of eurypterid from the Devonian of 
Gaspé. Proceedings of the Royal Society of Canada, 
Third Series, 46 (Appendix C): 149-150. [Abstract]. 
Cretaceous mammals of Alberta. Annual Report of 
the National Museum of Canada for the fiscal year 
1950-51, Bulletin 126: 110-119, Plates 14-15. 
Molluscan fauna of the Kishenehn Formation, South- 
eastern British Columbia. Annual Report of the Nation- 
al Museum of Canada for the fiscal year 1950-51, 
Bulletin 126: 120-141, Plates 16-19. 

Television and museums - second report. Bulletin of 
the Canadian Museums Association 5(4): 14-16. 
Gettysburg, a museum of fields and hillsides. Bulletin 
of the Canadian Museums Association 6(1): 10-12. 
Upper Cretaceous stratigraphy of southwestern Saskat- 
chewan. Billings Geological Society, Guidebook, 
Fourth Annual Field Conference: 87-97. 

Tertiary stratigraphy of southwestern Saskatchewan. 
Billings Geological Society, Guidebook, Fourth Annu- 
al Field Conference: 106-113. 

Museum news bulletins. Bulletin of the Canadian 
Museums Association 6(2): 8-11. 

Fauna and age of the Kishenehn Formation, south- 
eastern British Columbia. Proceedings of the Royal 


1953f 
1953¢ 


1953h 


1954b 


1954c 


1954d 


1954e 


1954f 


1955a 
1955b 
1955c 
1955d 


1955e 


1955f 


1955g 


1956a 


1956b 


1956c 


1956d 


1956e 


1956f 


461 


Society of Canada, Third Series, 47 (Appendix C): 
155. [Abstract]. 

The new art gallery of Hamilton. Bulletin of the 
Canadian Museums Association 6(4): 4. 

Museums of applied science. Bulletin of the Canadian 
Museums Association 6(4): 5-7. 

The new Kwakiutl house of Thunderbird Park. Bulletin 
of the Canadian Museums Association 6(4): 7-8. 


a A new species of eurypterid from the Devonian of 


Gaspé. Annual Report of the National Museum of 
Canada for the fiscal year 1952-53, Bulletin 132: 
83-91, Plates 1-2. 

Mammalian fauna of the Kishenehn Formation, 
southeastern British Columbia. Annual Report of 
the National Museum of Canada for the fiscal year 
1952-53, Bulletin 132: 92-111, Plates 1-3. 

Evidence of tooth structure on the relationships of 
the early groups of Carnivora. Evolution 8: 166-171. 
The Eocene-Oligocene transition as a time of major 
orogeny in Western North America. Proceedings of the 
Royal Society of Canada, Third Series, 48 (Appendix 
C): 35. [Abstract]. 

The Eocene-Oligocene transition as a time of major 
orogeny in Western North America. Transactions of 
the Royal Society of Canada, Third Series, 48 (Sec- 
tion 4): 65-69. 

A new species of Cephalaspis from the Devonian 
Gaspé sandstone at D’ Aiguillon. Le Naturaliste Cana- 
dien 81: 245-254. 

Dutch painting — the golden age. Bulletin of the Cana- 
dian Museums Association 8(1): 1-3. 

Television and museums — third report. Bulletin of 
the Canadian Museums Association 8(1): 10-16. 
Opening of the new Saskatchewan museum. Bulletin 
of the Canadian Museums Association 8(2): 8. 

Fort Ticonderoga bicentenary. Bulletin of the Canadian 
Museums Association 8(2): 11-13. 

Age of the Princeton Group, southwestern British 
Columbia. Proceedings of the Royal Society of Canada, 
Third Series, 49 (Appendix C): 40. [Abstract]. 
Recollections of some Alberta museums. Bulletin of 
the Canadian Museums Association 8(3): 9-12. 
Additions to the molluscan fauna of the Kishenehn 
Formation, Southeastern British Columbia and adja- 
cent Montana. Annual Report of the National Muse- 
um of Canada for the fiscal year 1953-54, Bulletin 
136: 102-119, Plates 1-3. 

Plastics in the museum. Bulletin of the Canadian 
Museums Association 9(1): 10-15. 

Additional occurrences of fossil horse remains in 
western Canada. Annual Report of the National Muse- 
um of Canada for the fiscal year 1954-55, Bulletin 
142: 153-154. 

New discoveries of Miocene vertebrates in Saskatche- 
wan. Proceedings of the Royal Society of Canada, 
Third Series, 50 (Appendix C): 39. [Abstract]. 
Nonmarine Mollusca from the North Park Formation 
of Saratoga Valley, Wyoming. Journal of Paleontology 
30: 1260-1263. 

The National Museum of History, Mexico. Bulletin 
of the Canadian Museums Association 9(4): 15-16. 
The Cretaceous reptile Champsosaurus natator Parks. 
National Museum of Canada Bulletin 145: 1-51, 
Plates 1-12. 


462 


1957a 


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1957c 


1957d 


1957e 


1957f 


1958a 


1958b 


1958c 


1958d 


1958e 


1958f 


1959a 


1959b 


1959¢c 


1959d 


1960a 


1960b 


196la 


1961b 


THE CANADIAN FIELD-NATURALIST 


Tenth anniversary of the Canadian Museums Asso- 
ciation. Bulletin of the Canadian Museums Associa- 
tion 10(1): 1-2. 

International Geophysical Year exhibit at the National 
Museum. Bulletin of the Canadian Museums Asso- 
ciation 10(1): 8-12. 

One man’s impressions of the Lincoln Nebraska meet- 
ing of the American Association of Museums. Bulletin 
of the Canadian Museums Association 10(2): 10-14. 
The eleventh annual meeting of the Northeast Muse- 
ums Conference, Montreal and Quebec cities, Sep- 
tember 3rd, 4th and Sth, 1957. Bulletin of the Cana- 
dian Museums Association 10(3): 1-5. 

Historical conservation along the St. Lawrence Sea- 
way. Bulletin of the Canadian Museums Association 
10(3): 7-9. 

Tertiary plains of Alberta and Saskatchewan. Proceed- 
ings of the Geological Association of Canada 9: 17-19. 
Mollusca from the Tertiary of Princeton, British 
Columbia. Annual Report of the National Museum 
of Canada for the fiscal year 1955-56, Bulletin 147: 
84-95, Plates 1-2. 

Paleocene mammal teeth from Alberta. Annual Report 
of the National Museum of Canada for the fiscal 
year 1955-56, Bulletin 147: 96-103, Plate 1. 

Report on the annual meeting of the Canadian Muse- 
ums Association, Windsor, Ontario, May 7th to 9th, 
1958. Bulletin of the Canadian Museums Associa- 
tion 11(2): 3-15. 

Mammal teeth from the Edmonton Formation at Scab- 
by Butte, Alberta. Proceedings of the Royal Society of 
Canada, Third Series, 52 (Appendix C): 24. [Abstract]. 
A horse astragalus from the Hand Hills conglomer- 
ate of Alberta. National Museum of Canada, Natural 
History Papers 1: 1-3. 

A palaeontological view of convergence, parallelism 
and orthogenesis. XVth International Congress of 
Zoology, Section 1, Paper 24. [Abstract]. 

The dentition of rabbits and the origin of the Lago- 
morpha. National Museum of Canada Contributions 
to Zoology for 1958, Bulletin 166: 41-45. 
Continental zoology of the North American Pleis- 
tocene. Problems of the Pleistocene and Arctic, Pub- 
lications of the McGill University Museums 1(1): 
39-45. 

Fossil mammals and intercontinental connections. 
Proceedings of the Royal Society of Canada, Third 
Series, 52 (Appendix C): 18. [Abstract]. 

Darwin: the origin of species. Pamphlet to accompa- 
ny an exhibit at the National Museum of Canada 
celebrating the 100th anniversary of the publication 
of “The Origin of Species”. 3 pages. 

The geological record of evolution. Pages 3-11 in 
Evolution: Its Science and Doctrine. Edited by T. W. 
M. Cameron. University of Toronto Press, Toronto. 
Fossil mammals and intercontinental connections. 
Pages 63-78 in Evolution: Its Science and Doctrine. 
Edited by T. W. M. Cameron. University of Toronto 
Press, Toronto. 

The National Museum of Canada 1910 to 1960. 
Department of Northern Affairs and National Re- 
sources, Ottawa. 37 pages. 

A museum for Canadians. Annual Report for 1960- 
61, Department of Northern Affairs and National 
Resources, 13 pages. 


1961c 


1962a 


1962b 


1962c 
1963a 
1963b 
1964a 
1964b 
1964c 
1965a 
1965b 
1965c 
1965d 


1965e 


1965f 


1965¢ 


1965h 


1965i 


1965} 


1966a 
1966b 


1966c 


1966d 


1966e 


1966f 


Vol. 118 


Swinton to ROM. Bulletin of the Canadian Muse- 
ums Association 13(4): 5. 

Mammalian migrations in the Pleistocene. Problems 
of the Pleistocene and Arctic, Publications of the 
McGill University Museums 2(2): 48-55. 

Mammal teeth from the St. Mary River Formation 
(Upper Cretaceous) at Scabby Butte, Alberta. Nation- 
al Museum of Canada, Natural History Papers 14: 1-4. 
Rudolph Martin Anderson. Canadian Geographical 
Journal 45(6): 198-199. 

Problems and potentialities of the history museum. 
Curator 6: 341-349. 

Canadian museums — this centenary and the next. Bul- 
letin of the Canadian Museum Association 14(4): 8-14. 
Kishenehn Formation. Bulletin of Canadian Petroleum 
Geology 12: 536-543. 

Cretaceous non-marine faunas of northwestern North 
America. Royal Ontario Museum Life Sciences 
Contribution 61: 1-24. 

The coming of kerosene. The Rushlight 30(2): 3-6. 
The problem of the Willow Creek Formation. Cana- 
dian Journal of Earth Sciences 2: 11-14. 

Pushing back the dawn. Meeting Place, Journal of 
the Royal Ontario Museum 1(3): 82-87. 

Body temperature of dinosaurs and its relationships to 
their extinction. Journal of Paleontology 39: 497-501. 
Alice Evelyn Wilson, 1881-1964. Canadian Field- 
Naturalist 79: 159-161. 

Frank Harris McLearn, 1885-1964. Proceedings of 
the Royal Society of Canada, Fourth Series, 3: 135- 
139. 

The mastodon. Royal Ontario Museum “What? Why? 
When? How? Where? Who?” 6: 1-16. 

Tertiary mammals of Saskatchewan, Part I: The 
Eocene fauna. Royal Ontario Museum Life Sciences 
Contribution 67: 1-33, Plates 1-7. 

W. A. Clemens and L. S. Russell. Mammalian fossils 
from the Upper Edmonton Formation. Pages 32-40 
in Vertebrate Palaeontology in Alberta. Report of a 
Conference held at the University of Alberta, Aug. 
29-Sept. 3, 1963. University of Alberta, Edmonton. 
The continental Tertiary of Western Canada. Pages 
41-52 in Vertebrate Palaeontology in Alberta. Report of 
a conference held at the University of Alberta Aug. 29 
to Sept. 3, 1963. University of Alberta, Edmonton. 
Macropalaeontology of the surface formations, Cypress 
Hills area, Alberta and Saskatchewan. Alberta Society 
of Petroleum Geologists, 15th Annual Field Conference 
Guide Book, Part 1, Cypress Hills Plateau: 131-136. 
A Paleocene conglomerate in westcentral Alberta. 
Canadian Journal of Earth Sciences 3: 127-128. 
Dinosaur hunting in western Canada. Royal Ontario 
Museum Life Sciences Contribution 70: 1-37. 

The changing environment of the dinosaurs in North 
America. The Advancement of Science, 23(110): 
197-204. 

Palaeontology of the Swan Hills area, north-central 
Alberta. Transactions of the Royal Society of Canada, 
Fourth Series, 4 (Appendix): 16. [Abstract]. 
Exploring the “New Red Sandstone”. Meeting Place, 
Journal of the Royal Ontario Museum 1(8): 105-107. 
Lighting the pioneer Ontario home. Royal Ontario 
Museum “What? Why? When? How? Where? Who?” 
12: 1-16. 


2004 


1966g 
1967a 


1967b 


1967c 


19674 
1967e 
1967f 
1967g 


1968a 


1968b 


1968c 
1969a 


1969b 
1970a 


1970b 


1970c 
| 1971 

1972a 
1972b 


| 1972c 
| 1972d 


—-1973a 


| 1973b 


1973c 


| 1974a 


1974b 


SEYMOUR: TRIBUTE TO LORIS SHANO RUSSELL 


More heat than light. The Rushlight 32(2): 1-4. 
Palaeontology of the Swan Hills area, north-central 
Alberta. Royal Ontario Museum Life Sciences Con- 
tribution 71: 1-31, Plate 1. 

Comment on: The inability of dinosaurs to hibernate 
as a possible key factor in their extinction, by J. M. 
Cys. Journal of Paleontology 41: 267. 
Confederation lamps. Canadian Antiques Collector 
2(3): 9-11. 

Review of: Fossil mammals of the type Lance For- 
mation of Wyoming, Part I: Marsupialia by W. A. 
Clemens. Journal of Paleontology 41: 813-814. 
Review of: Marsh’s Dinosaurs: The collections from 
Como Bluffs by J.-H. Ostrom and J.S. McIntosh. 
Journal of Paleontology 41: 1029-1030. 

Unionidae from the Cretaceous and Tertiary of Alber- 
ta and Montana. Journal of Paleontology 41: 1116- 
1120. 

A pedunculate cirripede from Upper Cretaceous rocks 
of Saskatchewan. Journal of Paleontology 41: 1544- 
1547. 

A dinosaur bone from Willow Creek beds in Montana. 
Canadian Journal of Earth Sciences 5: 327-329. 

A new cetacean from Oligocene Sooke Formation of 
Vancouver Island. Canadian Journal of Earth Sci- 
ences 5: 929-933, Plates 1-2. 

A Heritage of Light. University of Toronto Press, 
Toronto. 344 pages. 

Banquet lamp from the 1870s — a correction. Ontario 
Showcase 34(4): 6-7. 

Adventures in old-time lighting. Rotunda 2(1): 16-25. 
The great Saskatchewan Mouse Mine. Rotunda 3(1): 
16-24. 

Can we neglect research? Museum News 48(6): 13- 
14, 48. 

Correlation of the Upper Cretaceous Montana Group 
between southern Alberta and Montana. Canadian 
Journal of Earth Sciences 7: 1099-1108. 

Those remarkable dinosaurs. Rotunda 4(1): 4-17. 

L. S. Russell and C. S. Churcher. Vertebrate palaeon- 
tology, Cretaceous to Recent, Interior Plains, Canada. 
XXIV International Geological Congress (Montreal, 
Canada), Field Excursion Guidebook 24, Part A21: 
1-46. 

Tertiary mammals of Saskatchewan, Part II: The 
Oligocene fauna, non-ungulate orders. Royal Ontario 
Museum Life Sciences Contributions 84: 1-97. 
Report of the President. Proceedings of the Royal 
Canadian Institute, Series 5, 19: 2-5. 

The fifty million year pedigree of the horse. Proceed- 
ings of the Royal Canadian Institute, Series 5, 19: 6-15. 
Geological evidence on the extinction of some large 
terrestrial vertebrates. Canadian Journal of Earth Sci- 
ences 10: 140-145. 

Everyday Life in Colonial Canada. Copp Clark Pub- 
lishing Company, Toronto. 207 pages. 

Erratum: Geological evidence on the extinction of 
some large terrestrial vertebrates. Canadian Journal 
of Earth Sciences 10: 1361. 

R. E. Sloan and L. S. Russell. Mammals from the 
St. Mary River Formation (Cretaceous) of southwest- 
ern Alberta. Royal Ontario Museum Life Sciences 
Contribution 95: 1-21. 

Fauna and correlation of the Ravenscrag Formation 
(Paleocene) of southwestern Saskatchewan. Royal 


1974c 


1974d 


1975a 


1975b 


1975c 


1975d 


1975e 


1976a 


1976b 


1976c 


1976d 


1976e 


1976f 


1977 


1978 


1979 


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1980b 


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Ontario Museum Life Sciences Contribution 102: 1- 
53). 

Tools of the Trades. Pages 227-239 in The Book of 
Canadian Antiques. Edited by D. B. Webster. McGraw- 
Hill Ryerson, Toronto. 

Review of: Cars of Canada by H. Durnford and G. 
Baechler. Canada: An Historical Magazine 2(1 ): 59-60. 
Lower Cretaceous non-marine fauna of Alberta and 
Montana. Geological Society of America, Abstracts 
with Programs 7(6): 847. [Abstract] 

Revision of the fossil horses from the Cypress Hills 
Formation (Lower Oligocene) of Saskatchewan. Cana- 
dian Journal of Earth Sciences 12: 636-648, Plates 1-6. 
Mammalian faunal sucession in the Cretaceous sys- 
tem of Western North America. Pages 137-161 in W. 
G. E. Caldwell (editor), The Cretaceous System in the 
Western Interior of North America. Geological Asso- 
ciation of Canada Special Paper 13. 

“Carbide” Willson. Canada: An Historical Magazine 
3(1): 20-33. 

The first Canadian cooking stove. Canada: An Histor- 
ical Magazine 3(2): 34-35. 

Pelecypods of the Hell Creek Formation (Uppermost 
Cretaceous) of Garfield County, Montana. Canadian 
Journal of Earth Sciences 13: 365-388, Plates 1-7. 
The image of palaeontology. Pages 14-17 in Athlon: 
Essays in Honour of Loris Shano Russell. Edited by 
C. S. Churcher. Royal Ontario Museum Life Sci- 
ences Miscellaneous Publication. 

The palaeogeographic significance of the polypro- 
todont marsupials. XX Vth International Geological 
Congress, Sydney, Australia, Abstracts 1(7C): 333- 
334. [Abstract]. 

A new species of talpid insectivore from the Miocene 
of Saskatchewan. Canadian Journal of Earth Sciences 
13: 1602-1607. 

Early nineteenth-century lighting. Pages 186-201 in 
Building early America: contributions toward a his- 
tory of a great industry. Edited by C. E. Peterson. 
Chilton Books Company, Radnor, Pennsylvania. 
Early Canadian sewing machines. Canadian Collector 
11(5): 26-29. 

L. S. Russell and G. Edmund. Obituary: Levi Stern- 
berg, 1894-1976. News Bulletin of the Society of 
Vertebrate Paleontology 110: 43-44. 

Tertiary mammals of Saskatchewan, Part IV: The 
Oligocene anthracotheres. Royal Ontario Museum 
Life Sciences Contributions 115: 1-16. 

Handy Things to Have Around the House. McGraw- 
Hill Ryerson Limited, Toronto. 176 pages. 

Tertiary mammals of Saskatchewan, Part V: The 
Oligocene entelodonts. Royal Ontario Museum Life 
Sciences Contributions 122: 1-42. 

A new species of Brachyhyops? (Mammalia, Artio- 
dactyla) from the Oligocene Cypress Hills Forma- 
tion of Saskatchewan. Royal Ontario Museum Life 
Sciences Occasional Paper 33: 1-8. 

Alf Erling Porsild 1901-1977. Proceedings of the 
Royal Society of Canada, Fourth Series, 18: 111-114. 
Invention and discovery/Invention et découverte: the 
role of the inventor in Canada and the use of the patent 
office records. Pages 120-128 in Science, Technology, 
and Canadian History. Edited by R. A. Jarrell and N. 
R. Ball. Wilfrid Laurier University Press, Waterloo, 
Ontario. 


464 


1982a 


1982b 
1982c 


1982d 


1983 


1984a 


1984b 


1986 


1987a 


1987b 


THE CANADIAN FIELD-NATURALIST 


Tertiary mammals of Saskatchewan, Part VI: The 
Oligocene rhinoceroses. Royal Ontario Museum Life 
Sciences Contributions 133: 1-58. 

Charles Mortram Sternberg, 1885-1981. News Bulletin 
of the Society of Vertebrate Paleontology 124: 70-72. 
Charles Mortram Sternberg, 1885-1981. Canadian 
Field-Naturalist 96: 483-486, and cover photograph. 
Charles Mortram Sternberg, 1885-1981. Proceed- 
ings of the Royal Society of Canada, Fourth Series, 
20: 132-135. 

Evidence for an unconformity at the Scollard-Battle 
contact, Upper Cretaceous strata, Alberta. Canadian 
Journal of Earth Sciences 20: 1219-1231. 

Tertiary mammals of Saskatchewan, Part VII: Oligo- 
cene marsupials. Royal Ontario Museum Life Sci- 
ences Contributions 139: 1-13. 

Bringing them back alive. Rotunda 17(2): 12-17. 
Exploring a great dinosaur graveyard. Rotunda 19(2): 
20-29. 

Biostratigraphy and palaeontology of the Scollard 
Formation, Late Cretaceous and Paleocene of Alberta. 
Royal Ontario Museum Life Sciences Contributions 
147: 1-23. 

R. J. Emry, L. S. Russell and P. R. Bjork. The 
Chadronian, Orellan, and Whitneyan North American 


1988 
1989 


1990a 


1990b 


1993 


2003 


Vol. 118 


land mammal ages. Pages 118-152 in Cenozoic Mam- 
mals of North America: geochronology and _bios- 
tratigraphy. Edited by M. O. Woodburne. University 
of California Press, Berkeley. 

The first fossil hunters. Alberta 1(1): 11-16. 

Preface. Pages 1-2 in Geological History of Saskatche- 
wan by J. Storer. Saskatchewan Museum of Natural 
History, Regina. 90 pages. 

Thomas Chesmer Weston and the Red Deer River fossil 
fields. Earth Sciences History 9(1): 3-5. 

Charles Mortram Sternberg and the Alberta dinosaurs. 
Pages 1x-xill in Dinosaur Systematics: Approaches 
and Perspectives. Edited by K. Carpenter and P. J. 
Currie. Cambridge University Press, Cambridge. 

D. F. Stott, J. Dixon, J. R. Dietrich, D. H. McNeil, 
L. S. Russell, and A. R. Sweet. Tertiary. Pages 439- 
465 in Sedimentary Cover of the Craton in Canada. 
Edited by D. F. Stott and J. D. Aiken. Geological 
Survey of Canada, Geology of Canada 5. 

A Heritage of Light. University of Toronto Press, 
Toronto. 344 pages. [reprint of 1968 volume]. 


unpublished Biostratigraphy of the Horseshoe Canyon For- 


mation, Edmonton Group, Upper Cretaceous of Alber- 
ta. Unpublished manuscript on file at ROM. 


Book Reviews 


ZOOLOGY 


The Uses and Curation of Birds’ Egg Collections: An Examination and Biography 


By Martin Limbert. Peregrine Books, 27 Hunger Hills Avenue, 
Horsforth, Leeds, West Yorkshire LS18 S5JS. Hardcover. 
97 pages, £31 which includes postage to Canada if paid 
by postal money order. 


This book was written largely to explain why egg 
collections should be preserved and to counter nega- 
tive attitudes in Britain, where egg collecting has been 
illegal for half a century. Sadly, some museums have 
progressively removed birds’ eggs from public display 
and consigned them instead to storage and, often, neg- 
lect. The need for and methods of careful preservation 
of surviving egg collections are discussed in some 
detail. 

There is much here of historical interest. System- 
atic egg collecting in England dates back at least to 
1662. Hewitson in 1831 coined the term, “oology.” 
Overzealous collecting at times contributed to species’ 
declines. Egg set data are valuable in mapping distri- 
bution limits of individual bird species, but errors in 
identification and outright fraud by some egg dealers 
mean that some cannot be accepted at face value; the 
Canadian example of Walter Raine is mentioned in 
this regard. 

Limbert points out that nests and eggs have yet to 
be described for up to three thousand bird species in 
the world, and much information from extant collec- 
tions has not yet been shared. Only recently, numerous 
egg collections were studied to analyse 12 000 clutch- 
es of the Common Cuckoo, thereby adding greatly to 
our understanding of the extent of variation in size 
and colour of cuckoo eggs deposited in the nests of 
different hosts. Phenologic investigation reveals the 
extent to which the eggs are laid earlier in warmer 


Birds of Nunavut: A Checklist 


By James M. Richards, Y. Robert Tymstra, and Anthony W. 
White. Birders Journal, 701 Rossland Road East, Suite 
393, Whitby, Ontario LIN 9K3 Canada 21 pages. Canadian 
$8. Paper plus $1.50 postage. 


This attractive booklet lists 254 confirmed species, 
including 124 that are known to breed, plus 9 hypo- 
thetical species. Separate codes in columns give status 
codes for abundance and breeding for three arbitrary 
geographic subdivisions, “Mainland” (most but not all 
of the former territory of Keewatin), “Arctic Islands,” 
and “Bay Islands” (south of sixty degrees). As evi- 
dence that birding knowledge of Nunavut is increas- 


years. Eggs are occasionally useful in determining 
phylogenetic relationships of unusual species such as 
the Sandgrouse. Preserved contents of eggs can be 
saved for pesticide analysis. And, as modern ornithol- 
ogists learned from Ratcliffe in 1962, egg-shell thin- 
ning has been an indicator of pesticide burdens and 
thus a method of monitoring the environment. 

Of interest to many readers will be the gastronomic 
aspects of seabird egg-collecting on cliffs, especially 
the abundant and highly palatable egg of the Common 
Guillemot at Flamborough. Later, eggs from this guille- 
mot colony became celebrated among oologists for 
their variety of marking and colour. 

This book deals almost exclusively with oological 
matters — and oology publications — concerning Great 
Britain. Thirty-three valuable pages of bibliography 
are given, citing mainly European sources; the land- 
mark American publications of Bent, Bendire and 
Reed are not included. Oology periodicals began forty 
years later in Britain than those in the United States, 
but ceased publication about the same time. as the 
Oologist, the last survivor in America. 

I detected one error in terminology, the use of “ex- 
tinction” from one country of a species surviving else- 
where; he should have used “extirpation.” This attrac- 
tive, but expensive, little book is recommended to 
anyone interested in the British approach to past bird’s 
egg collections. 

C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N 0J8 
Canada 


ing rapidly, 48 confirmed species have been added 
during the three years immediately prior to publica- 
tion. An excellent map is unusually helpful because it 
gives both the former English names and the Inuktitut 
names (new for most of us) for each settlement, many 
of which were initially Hudson’s Bay Company fur 
trading posts. There are two pages of bibliography. All 
in all, this is a commendable work in progress, and 
the price is within easy reach. 

C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N 0J8 
Canada 


465 


466 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Birds of the Untamed West: The History of Birdlife in Nebraska, 1750 to 1875 


By James E. Ducey. 2000. Making History, [but released for 
review in 2002]. 2415 N 56th Street, Omaha, Nebraska 
68104 USA. 300 pages. U.S. $25.00. Paper. 


This book is a valiant attempt to report the ornitho- 
logical history of Nebraska to 1875. Commendable 
strengths include Chapter 1, which discusses the bird 
knowledge and lore of the native Americans, the Lako- 
ta, Missouria, Otoe, Omaha, Pawnee, Ponca, and Win- 
nebago tribes. Native language bird names are pro- 
vided when available. Chapter 2 provides a summary 
of historic explorations, most of which were made by 
men merely passing through the state while heading 
farther north and west. Many of these explorers came 
through in autumn, after the bird breeding season was 
over. Exceptions were Lewis and Clark, in Nebraska 
from 11 July to 8 September 1804, and Thomas Say 
with the Major Long expedition, present from 19 Sep- 
tember 1819 to 6 June 1820. Chapter 3 provides a suc- 
cinct account of the early bird habitats, and Chapter 4 
tells which species were found in each of these habi- 
tats. Ducey provides, in square brackets, occasional 
corrections of obviously misleading statements in 
Aughey’s 1877 paper. The list of references I found 
impressive. Thirty-three early illustrations add to the 
interest and attractiveness of the book. 

For each observation, the name of the current Ne- 
braska county is provided in upper case letters: “shout- 
ing” in modern computer parlance and to me a bit 
annoying. Unlike Robert E. Stewart’s North Dakota 
book, Ducey does not provide a map showing the loca- 
tion of each county, forcing the reader to provide him- 
self with a Nebraska state map before reading very 
far. 

Before listing the shortcomings of Chapter 5, the last 
half of the book, I chose to use Myron Swenk’s histori- 
cal articles in Nebraska Bird Review (in the late 1930s) 


as a veracity check. I was not too surprised to find that 
Swenk had, in the late 1930s, provided more detail and 
better documentation than Ducey. If one takes the Lewis 
and Clark expedition as an example, Swenk used a not 
excessive seven pages, including a map showing the 
progress day by day, consulted original, primary sources, 
and gave more detail about extant diaries of several 
members of the expedition. As a further check, I com- 
pared the four-plus pages of Nebraska citations in Gol- 
lop’s Eskimo Curlew monograph with Ducey’s account, 
which again was less complete. 

Chapter 5, a List of Species, occupies 110 pages; it 
lists excavated faunal remains from various forts and 
Indian camps, and is a useful compilation that leads 
the reader to original sources. Sadly, Ducey fails to 
place the verbatim accounts of each species in the ex- 
plorer’s words, indicated by quotation marks or a dif- 
ferent font. As a result, one can rarely differentiate fact 
from Ducey’s speculation, extrapolation, and “best 
guesses.” His terminology and presentation are incon- 
sistent, especially concerning whether an individual 
species 1s a migrant or a resident, and whether or not 
there is specific evidence of breeding. His use of “migra- 
tory species” is a less satisfactory term than “migrant.” 
Far too often, the term “potential breeder” is used with- 
out evidence of dates or localities for eggs or young. 
Clearly, a conventional publishing house would have 
provided the outside editorial assistance that this 
book lacks. The index is incomplete. 

In spite of my caveats, especially the idiosyncratic 
presentation of the species list, anyone interested in 
the history of ornithology in Nebraska will find much 
of interest in this inexpensive book. 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N 0J8 
Canada 


Birds of Nebraska: Their Distribution and Temporal Occurrence 


By Roger S. Sharpe, W. Ross Silcock, and Joel G. Jorgensen. 
2001. University of Nebraska Press, 233 North 8" Street, 
Lincoln, Nebraska 68588-0255 USA 520 pages. U.S. 
$69.95. Cloth. 


This book is a nearly ideal state bird book, a scholar- 
ly compilation of sightings and specimens, with careful 
assessment of questionable sight and specimen records. 
Subspecies receive detailed and precise attention. There 
is helpful information about the best spots to search 
for each species. Good use is made of data from 
Breeding Bird Surveys and Christmas Bird Counts, 
but only sporadic use is made of banding recoveries. 
The price is kept low by omitting paintings of each 
bird species, a feature of more sumptuous state books. 
Four maps are useful for locating counties, but one 
must consult the Gazeteer (which fails to include Pine 
Ridge) to locate specific towns, parks, refuges and 
other geographic features. 


The introduction deals with geography, geomorphol- 
ogy, aquatic systems, and climate, and then describes 
each of the main environments. The history of Nebraska 
ornithology is given in ten succinct pages, while anoth- 
er six tell how to use the species accounts, emphasiz- 
ing the key importance of the Distribution and Ecolo- 
gy section for each species; sadly, the reader is not 
alerted to the list of abbreviations in Appendix 1. 

As might be expected in such a book, there is a 
wealth of intriguing information, especially about range 
extensions and both increases and decreases in popu- 
lations; commendably, the maximum number of indi- 
viduals seen at one time is provided for most species. 
Other items that caught my attention were: Gray Par- 
tridge spread into Nebraska spontaneously from South 
Dakota and Iowa after much earlier attempts to intro- 
duce them directly had failed; most Ruffed Grouse 
disappeared by the 1880s, and it is now listed as an 


2004 


extirpated species; a Clapper Rail caught in a mink trap 
in January was the farthest inland record, the nearest 
being Tennessee; several hundred Buff-breasted Sand- 
pipers followed a farmer working a field; a Cliff Swal- 
low colony contained 3700 nests. There is a useful 
warning about reliance on vocalizations to separate the 
Eastern from the Western Wood-Pewee. The authors 
also mention a population of chickadees in the Wild- 
cat Hills that resembled Black-capped Chickadees 
morphologically, but sang Mountain Chickadee songs. 

Regrettably, lower priority is given to nest records as 
compared to sightings, although the latter are of less- 
er biological importance. Only a relatively few species 
have data from the Cornell Nest Card Program, with 
detailed numbers and dates of nests. For some other 
breeding species, dates and localities of nests, eggs or 
flightless young are not provided; we can only guess 
that in some cases they were not available, but surely 
for at least the Mourning Dove and Yellow-headed 
Blackbird they were simply omitted. No list of host 
species for Nebraska is provided for the Brown-head- 
ed Cowbird. 

It would have been helpful to place unaccepted 
records within square brackets and add abbreviations 
for journals at the top of the list of references. The 


Book REVIEWS 


467 


subspecies account for the Great Horned Owl is incor- 
rect since the wapacuthu race of the Great Horned 
Owl has been discredited. The account of the Poor- 
will is misleading, implying that the Flint Hills are in 
Nebraska, whereas they are in Kansas. Mention is 
made of a Golden Eagle nest with four young with- 
out comment that this may be the first such record in 
the North American literature, though observations 
of two Golden Eagle nests with four eggs have been 
published previously. 

My minor criticisms aside, this is an excellent sum- 
mary of what is known about Nebraska birds. The state 
is important for the number of its bird species (415 
with a specimen or recognizable photograph, 13 with 
a description acceptable to the state committee, and 5 
extinct or extirpated), and because it is the meeting 
place for many closely related species which overlap 
in range and sometimes interbreed. Birds of Nebraska 
is a substantial and welcome addition to ornithology 
in general and to state bird books in particular. 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N OJ8 
Canada 


Canadian Skin and Scales: A Complete Encyclopedia of Canadian Amphibians and Reptiles 


By Pat E. Bumstead. Illustrated by Norman H. Worsley. 
2003. Simply Wild Publications Inc., 100 Lake Lucerne 
Close SE, Calgary. Alberta T2J 3H8 Canada 161 pages. 
$24.95. 


Although subtitled “a complete encyclopedia” this 
book obviously was never intended to be so preten- 
tious, but instead to serve young naturalists’ as a “first 
book”. The cover, with bright colour photos of a 
Horned Lizard (Stephen Glendining), Plains Garter 
Snake, Western Painted Turtle, Northern Leopard Frog, 
and Tiger Salamander (Brian Woltski), will attract 
them. Any suspected western bias is soon dispelled 
by the content, which is spread evenly across Canada. 
Black-and-white sketches by Norman H. Worsley de- 
pict all Canadian species and a selection of extra-limital 
representatives of families. Many are adequate, some 
good, but others are barely recognizable (e.g., the wrin- 
kled Spotted Salamander on page 25 and Black Rat 
Snake on page 124), and one is clearly the wrong 
selection (the Ensatina on page 32 of a southern pat- 
tern not occurring in Canada). 

The pedagogical style begins with “Canadian Crea- 
tures”, covering communal activities, wintering, being 
dark, food resources and hibernating sites, and advan- 
tages for those reptiles bearing young (in Canada, two 
lizards and 13 snakes) rather than laying eggs. 
Province-by-province species lists follow. Chapters 
2-9 deal with amphibians in general, salamanders and 
newts (21 species), frogs and toads (25), reptiles in 
general, turtles and tortoises (14), lizards (5), and snakes 


(24). Each family and each species which occurs in 
Canada is an individual account. Those for families 
give world and Canadian species totals, world range, 
characteristics, and reproduction. The species accounts 
include common and scientific name, a paragraph of 
introduction, colour, reproduction, “where do they 
live” (habitat and provinces), and a “did you know” 
section. A series of questions with reference to the 
page where an answer can be found are at the bottom 
of many of the pages in this section. Chapter 9 deals 
with Conservation and includes definitions for status 
categories and species thought to be extirpated in 
Canada (Timber Rattlesnake, Pacific Pond Turtle, 
Pygmy Horned Lizard, Pacific Gopher Snake, East- 
ern Tiger Salamander). Nine threats to others are List- 
ed from habitat loss to introduction of exotic species. 
A “you can help” section presents ideas for group and 
individual participation. A final chapter, “Etcetera”, 
covers diagrams of forms and features, the availability 
of a teachers guide, and a list of Canadian internet 
wildlife links, words to know, published and internet 
resources used, and a five-page index. Finally, there 
is a note from the author and a short biography of the 
illustrator. 

Although primarily concerned with Canada, the text 
erratically adds extra-limital filler statements such as 
that Mudpuppies “have been introduced in large New 
England rivers” or that Western Skinks have been found 
on islands off the coast of California. For the geograph- 
ically challenged it is not made clear that species listed 
for “Newfoundland” are those for the political entity 


468 


that includes mainland Labrador (accounting for the 
salamanders included), or that the generally arctic 
Nunavut extends south to include islands in Hudson 
Bay (accounting for the occurrence of a frog and a 
toad in the territory). 

The foreword, an endorsement by Carolyn and 
David Seburn, rightly extols the book for its overrid- 
ing themes of excitement in observing amphibians 
and reptiles, the need to treat them with respect, and 
the importance of conserving their habitats. But they 
overlook, or were unaware of, problems precipitated 
by generalizations and simplifications while copying 
information from the literature apparently without per- 
sonal experience with many forms. Particularly mis- 
leading in all frog and toad accounts is that the num- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


ber of eggs is followed by an “adults appear” which 
actually refers to when tadpoles transform, the result- 
ing froglets are not “adults” (mature) for months or 
another year or more later. For all toads, spadefoots, 
and treefrogs only the aquatic habitats where they 
breed are given under “where they live” whereas 
most are terrestrial much of the year. Unfortunately, 
such “information” is as easily absorbed by the un- 
wary and uncritical beginner as fact, and detracts from 
the otherwise commendable concept and aim of the 
effort. 


FRANCIS R. COOK 


Canadian Museum of Nature, Ottawa, Ontario K1P 6P4 
Canada 


Conservation and Ecology of Turtles of the Mid-Atlantic Region: A Symposium 


Edited by Christopher W. Swarth, Willem M. Roosenburg 
and Erik Kiviat. 2004. Bibliomania! books @biblioma 
nia.com. 122 pages. U.S. $22.50. 


The Mid-Atlantic region of the USA (from Vir- 
ginia to New York) is an area of exceptional turtle 
diversity, with 22 species (including four sea turtles). 
It is also an area under exceptional development 
pressure. A two-day conference was organized to dis- 
cuss the status and ecology of the species affected 
and held in October 1999. 

This volume brings together 11 peer-reviewed 
papers and 18 abstracts from the conference. The 
book begins with an introduction by the editors and the 
text of a keynote address by Michael Klemens, who 
briefly summarizes the conclusions from his book 
Turtle Conservation (2000; Smithsonian Institution 
Press). The papers cover only six of the possible species 
occurring in the area, with three papers on each of the 
Diamondback Terrapin and the Box Turtle, two on 
the Red-bellied Turtle and one each on the Blanding’s 
Turtle, Bog Turtle, and Spotted Turtle. The papers 
cover a wide range of topics including nest predation, 
head-starting, habitat change detection, and population 


For Love of Insects 


By Thomas Eisner. 2003. The Belknap Press of Harvard Uni- 
versity Press. Cambridge, Massachusetts, and London, 
England. 464 pages. U.S. $29.95. Cloth. 


Thomas Eisner is an entomological legend. His 
photo, on the dust jacket of this fine book, shows a 
middle-aged man cockily riding his bicycle, seated 
backward on the handle bars. Eisner is to entomology 
what Richard Feynman was to physics — brilliant, 
quirky, and full of good stories. If, for some reason, 
you need to be convinced of the fact that insects are 
among the most amazing creatures on earth, this is 
the book for you. 

The preface to this book of insect tales compares 
Thomas Eisner to Jean-Henri Fabre, the pioneer writer 
on insect behaviour, who lived in the 19" century in the 


ecology. Although the papers are peer-reviewed, they 
are of varying quality. One of the papers is barely more 
than a page in length and is little more than a report on 
the number of turtles caught at one site. From a con- 
servation perspective, the most interesting paper is by 
Erik Kiviat (one of the editors) and various collabo- 
rators and deals with the response of Blanding’s Turtles 
to wetland and upland habitat creation as part of a 
wetland mitigation project. Although the results are still 
preliminary (three years) Blanding’s Turtles made use 
of constructed nesting sites and wetlands. It is inter- 
esting, however, that the turtles did not choose to over- 
winter in constructed wetlands. 

This collection is not the definitive statement on 
the conservation of turtles in the eastern U.S. There are 
no papers (although some abstracts) on many topics, 
such as traffic mortality, or the effects of toxins, or 
genetic isolation. Nonetheless, it is a valuable snapshot 
of the wide range of activities being undertaken and it 
will be of interest to anyone involved in turtle conservation. 


DAVID SEBURN 


Seburn Ecological Services, 920 Mussell Road, RR 1, Oxford 
Mills, Ontario KOG 1S0 Canada 


south of France. E. O. Wilson, the preface’s author, 
seems comfortable with this comparison, but to me 
they are two very different sorts of scientists. Fabre 
was a poor man, and a loner. His observations were 
conducted with no institutional support, and his genius 
(Darwin called him “the incomparable observer”) was 
not recognized until Fabre was a very old man. Eis- 
ner, by contrast, is a hot-shot researcher at the top of 
his game, at what is probably the finest university for 
insect studies in North America (Cornell, in Ithaca, 
New York), surrounded by cooperative peers, gradu- 
ate students, and lots of grant money. While Fabre’s 
stories tell of hardship and isolation, Eisner’s explore 
the life of a modern biologist in the publish-or-perish 
world of research science. (Publishing, by the way, 


2004 


seems to come as easily to Eisner as sneezing comes 
to most of the rest of us.) 

The greatest thing about Eisner, however, is that he 
keeps the passion alive, and dwells not on the institu- 
tional politics of science, but on his life-long fascina- 
tion with the creatures that he studies. Eisner is a 
chemical ecologist, and thus the book is largely about 
insects and the chemicals they produce. But don’t get 
the impression that it is technically difficult to under- 
stand. It begins with a chapter on bombardier beetles, 
and the amazing way that they spray boiling quinines 
out their butts, and direct them accurately into the 
faces of their enemies. This chapter, like the others, 
does a nifty little dance between the insects and their 
adaptations on the one hand, and the process of sci- 
entific discovery on the other. The rest of the book is 
just as spellbinding, and in it the reader is treated to 
such juicy tidbits as explanations of how living things 
can defend themselves with cyanide without acciden- 
tally committing suicide, along with a host of other 
marvelous insect adaptations, all skilfully elucidated 
by Eisner, his colleagues, and his students. The chap- 
ter on spider webs is wonderful. And if you think you 
understand insects and mimicry, this book will surely 


The Freshwater Fishes of Manitoba 

By Kenneth W. Stewart and Douglas A. Watkinson. 2004. 
University of Manitoba Press, 301 St. John’s College, 
University of Manitoba, Winnipeg, Manitoba, R3T 2M5. 
xvii + 276 pages. $29.95. Paper. 


This book is a delight to read and use. The layout 
is innovative and the text is in an exceptionally clear 
font and is well written. The book is dedicated to the 
late Dr. Ed J. Crossman of the Royal Ontario Museum 
and co-author of the book Freshwater Fishes of Canada. 

The freshwater fishes of Manitoba comprise 79 
native species, | re-introduced species after extirpation, 
10 introduced species, 2 artificial hybrids and 4 estu- 
arine species from the Hudson Bay coast. This is the 
third most diverse ichthyofauna in Canada after Ontario 
and Quebec. Fifteen species from waters outside, but 
neighbouring Manitoba, are included as they may 
eventually be discovered in the province. There are 
various introductory sections such as biogeography, 
geography, species diversity patterns, and summary 
sections like a glossary and a checklist, usually found 
in fish books. There is also an appendix which sum- 
marises fish distributions by watershed and a list of 
references. There is no index but the unique layout 
assists in navigating the pages. 

The presentation of the book is very attractive and 
easy to use. Each family account with its species has a 
unique colour which appears in text headings, scientif- 
ic and common names here and in tables elsewhere, 
and along the upper half of the outer page margin (out- 
lining the English and French names) which allows 
rapid flipping as a search mechanism. It is immedi- 


BooK REVIEWS 


469 


expose you to vast unexpected dimensions to this sup- 
posedly simple phenomenon. 

Thomas Eisner is also a superb photographer, and 
one of the other great strengths of this book lies in 
the pictures. He also uses clever illustrations to make 
his point, and is clearly the sort of person who is 
good at entertaining his undergraduate students while 
he teaches. All of this comes together masterfully, to 
create a very fine book indeed. Do I have any criti- 
cisms? Not really, although for a book about “insects” 
it contains a wealth of information on arachnids as 
well. I suppose Eisner and his publisher didn’t want 
to use the term “arthropods,” or the term “bugs” to 
get around this perennial problem. No—this is a superb 
book, and a book that naturalists at all levels will 
enjoy. At the weekly entomology luncheon at the 
University of Alberta, I found that many of my senior 
colleagues (very well-read and enthusiastic entomolo- 
gists!) were amazed by how much they learned from 
“For Love of Insects.” I enthusiastically agree, and 
recommend it heartily. 


JOHN H. ACORN 


Department of Renewable Resources University of Alberta, 
Edmonton, Alberta, TST 5L7 Canada 


ately obvious when one moves from one family to 
another in the text and quick searches for a particular 
group are facilitated. The scientific and English fami- 
ly name is at the top right and left of each page and 
also allows rapid flip searches. 

The series of habitat photographs in the geography 
section are excellent, and have descriptive comments. 
One, showing the lower Churchill River could be 
almost anywhere in the vast boreal forest except for 
that peculiar hazard to Manitoban freshwater ichthy- 
ologists, a Polar Bear paddling by. 

Each species account gives the English, French 
and scientific names, a colour photograph of the fish, 
a section on Identification with key characters in bold 
(sometimes only a single, short sentence for distinc- 
tive species), a Distribution in Manitoba, Biological 
Notes including spawning, growth and adult size, feed- 
ing, habitat, and ecological role, and Importance to 
People. The latter refers to commercial, angling, eco- 
logical and conservation importance. There is no 
lengthy anatomical description of the species as is seen 
in most fish books, characters being restricted to those 
used in identification with some explanation of colour 
variations and amplification of characters from the 
Keys. 

All the fish illustrations are ideally positioned in 
the species description rather than grouped as colour 
plates. These photographs are generally excellent, al- 
though some key characters such as mouth parts are 
not evident and a close-up photograph of them would 
have added to the reader’s understanding. 


470 


The distribution of fishes on the maps is very clear. 
Red spots show known occurrences, black spots intro- 
ductions while grey shading shows the continuous range 
of native species. The grey shading generally conforms 
to distributions based on the red spots, filling in the 
gaps. However in a number of species, the limits of grey 
shading conform neither to the red spots nor obvious- 
ly to drainage basins. There is no explanation of how 
this overall distribution was arrived at but is presum- 
ably related to ecological limits for the species and 
physical barriers. Some form of shading to fill in lakes 
would have made the background map clearer and 
perhaps partially address the previous comment. 

Identification keys for each family are found at the 
beginning of each family account. An alternative ar- 
rangement is to group all keys together and this is 
one many field and laboratory biologists would pre- 
fer. Fish are wet and slimy and having keys grouped 
together makes for less page turning or facilitates xerox- 
ing and annotating. The keys work well although the 
one for distinguishing the Brown and Black bullheads 
gives diagrams of the supraethmoid shapes without 
explaining what or where this structure is (and it is 
not in the Glossary either). 

Some other minor points of criticism and comment 
must be noted. The scientific names lack the author 
and date, perhaps not of significance to most readers 
but a nuisance to find for those not familiar with the 
ichthyological literature. The copy I have lacks paper 
covers and there is only the briefest of blurbs on the 
authors whose names are sufficiently common not to 
lend themselves to easy Googling — more background 
on the authors is often of interest to readers. The com- 
mon names of fishes in English and French are on the 
margin of each page enabling a rapid flip through as 
a search. However the scientific name is not there 
(and there is space to add it) which would have been 
a great convenience. It is necessary to remember to 
flip back to front as well as front to back since some 
species are only on one page; this is inescapable in 
the design. The key to Cottidae is a little confusing as 
“Key to the freshwater sculpins and marine sculpins 


The History of Ornithology in Virginia 


By David W. Johnston. 2003. The University of Virginia 
Press, Box 400318, Charlottesville, Virginia 22904-4318 
USA. 219 pages. $35.00 U.S. Cloth. 


True to the promise implicit in his title, David John- 
ston’s book is a thorough history of ornithology in 
Virginia. The first five chapters take us from the Ter- 
tiary period 65 million years ago through to an exam- 
ination of 19th-century reports and collections. Along 
the way we learn all manner of interesting facts. The 
abundance of skeletal elements of the extinct Passen- 
ger Pigeon show it to have been common in the late 
Pleistocene over 12,000 years ago, and Rock Ptarmi- 
gan, Spruce Grouse and Gray Jay remains indicate a 
cooler climate at the time. Archaeological research at 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


of the genus Myoxocephalus” as this can be read as 
being a key to only Myoxocephalus species although 
Cottus species and freshwater Myoxocephalus are 
included. The photographs note whether a specimen 
is fresh or preserved but readers should be warned that 
preserved here generally means very recent preserva- 
tion as colour is still retained — museum specimens 
of any age soon lose the colour; compare the colour- 
ful “preserved” northern redbelly dace with the really 
preserved and brownish deepwater sculpin. A few 
photographs are not as revealing as one could hope 
for, the colourful sticklebacks being poorly served in 
this respect. The Glossary is good, distinguishing such 
terms as bar, band and stripe although commensial is 
more commonly spelled commensal and watershed is 
not a “water body together with all its tributaries” but 
strictly “an elevated boundary area separating tribu- 
taries draining to different river systems”, and the 
subopercular bone does not lie completely above the 
interopercular but mainly behind it (see www.brian- 
coad.com). 

Although books on fishes do tend to separate into 
those on marine species and those in fresh waters, the 
Manitoban coastal fauna on Hudson Bay comprises 
only 27 species (Coad and Reist 2004) and could 
have been included in the book. The authors do in 
fact deal with 13 of these species which are found in 
estuarine and fresh waters also. So another 14 spe- 
cies would have given a complete treatment of the 
Manitoban fishes. 

This book will long stand as the definitive guide to 
Manitoban freshwater fishes and sets a standard for 
all subsequent provincial and national books of fish 
faunas. 


Literature Cited 

Coad, B. W., and Reist, J. D. 2004. Annotated List of the Arctic 
Marine Fishes of Canada. Canadian Manuscript Report 
of Fisheries and Aquatic Sciences, 2674: iv + 112 pages. 


BRIAN W. COAD 


Canadian Museum of Nature, PO Box 3443, Station Douglas, 
Ottawa, Ontario K1P 6P4 Canada 


a 1700-year-old Indian midden provided the state’s only 
record of the Ivory-billed Woodpecker. Around 1650, 
early English settlers were so hungry that at times 
they ate bluebirds, larks, cardinals and goldfinches — 
and Carolina Parakeets — as well as waterfowl, shore- 
birds, and upland game birds. When settlers arrived 
from England about 1590, the Indians were cutting 
holes in gourds to entice Purple Martins to nest. 

The chapters on early observers and naturalists are 
particularly interesting. Thomas Hariot, a young man 
selected by Sir Walter Raleigh to be part of the 1585 
expedition and settlement, made one of the first 
attempts to list North American birds, but Johnston 
tells us that Hariot’s list of 111 species has never been 
found. Fortunately, the paintings of John White, Har- 


2004 


iot’s fellow expedition member and grandfather of 
Virginia Dare (the first English child born in Ameri- 
ca), have survived. White painted 35 species of birds 
and gave the Algonquian name for 23 of them. 

The first naturalists in Virginia were the clergyman 
and botanist, John Banister, who lived at Bristol Parish 
from 1678 until he was accidentally killed in 1692, 
and the Reverend John Clayton (1657-1725), who 
published information about birds and weather in the 
world’s first scientific journal, Philosophical Transac- 
tions of the Royal Society of London; four pages of 
Clayton’s account are reproduced by Johnston. Sadly, 
although Mark Catesby spent seven years in Virginia, 
1712-1719, he was then a botanical collector who 
had not yet learned to paint birds; from those years 
he mentioned only 33 bird species. When he returned 
later for another six years, Catesby spent his time far- 
ther south, so that his famous book deals mainly with 
the Carolinas and rarely makes specific mention of 
Virginia. In 1787, Thomas Jefferson, 14 years before 
he became the third president of the United States, 
made the first attempt to list all the birds of Virginia, 
adding 34 additional species to the 100 pictured for 
adjacent regions by Catesby. 

Later chapters deal specifically with topics such as 
the contributions of ornithologists with the Smithson- 
ian Institution and the United States government; con- 
servation and game laws; artist-naturalists; extirpated 
and introduced bird species; falconry; and regions of 
ornithological importance describing the observers 


All-Weather Hawk Watcher’s Field Journal 


By Donald Heintzelman. J. L. Darling Corp. 2614 Pacific 
Hwy., Tacoma, Washington 98424-1017 USA. 66 pages. 
U.S. $7.95 Paper 


This handy pocket field notebook, 12 by 17.5 cm., 
consists of a conservation note, a one-page introduc- 
tion, a three-page list of the diurnal birds of prey of 
North and Central America, and 51 pages for field 
observations. At the top of each page are blanks to 


Self-Portrait With Turtles: A Memoir 


By David Carroll. 2004. Houghton Mifflin, 222 Berkeley 
Street, Boston, Massachusetts USA. 181 pages. U.S. $23. 


David Carroll first saw a Spotted Turtle at the age 
of eight. He has been enchanted by these amazing 
creatures ever since. 

In this exquisitely written book the author of The 
Year of the Turtle and Swampwalker’s Journal shares 
his obsession with turtles, nature and art. This basi- 
cally chronological book is divided into four sections: 
Early Years, Art School, Middle Years, Later Years. 
Through these sections we see the development of a 
consummate naturalist and artist, witness his choice of 
art school over science and the growth of his teaching 
and artistic careers. The book concludes with Carroll 
hunting for turtles on the 50" anniversary of his first 
discovery of a Spotted Turtle. 


BOOK REVIEWS 


47] 


and the contributions for each. Many famous natural- 
ists worked in or passed through Virginia, among 
them John James Audubon and Roger Tory Peterson. 
Peterson, while stationed with the U.S. army at Fort 
Belvoir, “successfully petitioned the officer-in-charge 
to reroute the line of march on the drill field to avoid 
an occupied Horned Lark’s nest” (page 121). 

Items deserving special commendation are the 
detailed lists: principal ornithological accomplishments; 
type specimens from Virginia; local bird lists, 1870- 
1926; recent bird lists since 1952; Virginia nature 
writing, 1817-1998; books since 1965 that mention 
Virginia’s avifauna; Algonquian Indian names of birds; 
bird banders, 1923-1965; principal collectors of bird 
specimens; and observers reporting migration records 
to the U.S. Biological Survey, 1884-1946. Detailed 
references throughout and a selection of old draw- 
ings and more recent photographs add to the interest. 

Johnston’s scholarly and painstaking research makes 
this is one of the finest ornithological histories avail- 
able for any state. There are some weaknesses, such 
as the lack of a Virginia map and an incomplete and 
inconsistent index which omits names of some impor- 
tant people. This book is a necessity in every museum 
and University library in North America, and for any 
one with an interest in the history of ornithology. 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N 0J8, 
Canada 


fill in for date, time, weather, and location, including 
GPS coordinates. The special feature is the use of all- 
weather writing paper so that one can write in the 
rain! 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N OJ8, 
Canada 


Carroll’s memoir overcomes the common pitfall of 
bogging down in autobiographical trivia. His writing 
is compelling and thought-provoking: “Consecrated 
to the God of my parents before my eyes were open, 
I lived my first eight years in a closed circle of family, 
relatives, church, and school. I lived in a totally human 
environment filled with human concerns and consid- 
erations. It was a world built by people for people.” 
And yet within three days of his family moving to a 
new home, Carroll had discovered a wetland and en- 
countered a Spotted Turtle: “With that first turtle I 
crossed a boundary of greater dimensions than I can 
ever fully comprehend. I changed lives within a life, 
worlds within a world.” 

Although Carroll’s passion is turtles, his deep con- 
nection with nature will resonate with any avid natu- 


472 


ralist. His writing is honest and moving in its evoca- 
tion of special places and moments as well as the loss 
of many of those special places over time: “My long 
history with turtles has been marked time and again 
by loss of place, by the physical and spiritual annihi- 
lation of the landscape, compelling me to move on in 
search of wilder places.” 

This is also the kind of book you can give to non- 
naturalist friends to try to make them understand why 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


you love wading through swamps. Its combination of 
graceful writing, compelling anecdotes and Carroll’s 
own beautiful black and white illustrations are enough 
to enchant almost any intelligent reader. 


DAVID SEBURN 


Seburn Ecological Services, 920 Mussell Road, RR 1, Oxford 
Mills, Ontario KOG 1SO Canada 


Wild Mammals of North America: Biology, Management, and Conservation (Second edition) 


Edited by George A. Feldhamer, Bruce C. Thompson, and 
Joseph A. Chapman. 2003. The Johns Hopkins University 
Press, 2715 North Charles Street, Baltimore, Maryland 
21218-4363 USA. xiii + 1216 pages. U.S. $175. Cloth. 


After a 21 year interlude, the update of this monu- 
mental volume on the biology and management of 
North American mammals has added a third editor 
and given a more prominent role to conservation as 
opposed to economic importance. Even with a larger 
page format and a reduction of two chapters, the sec- 
ond edition is longer than the first, which reflects the 
accumulated increase in research over the past two 
decades. All of the accounts are updated to various 
degrees with some references as recent as the same 
year of publication (2003) of this book. A completely 
new set of authors has been recruited to write half (28) 
of the 55 chapters. Only six chapters have retained the 
original contributors and these are all single authored 
accounts. However, three of these accounts (black bear, 
badger, and manatee) are negligibly changed from the 
first edition. Of the 102 authors, there are three who 
have contributed to two chapters. 

The species coverage of this revised volume has 
been slightly modified. There are new accounts for 
Cynomys ludovicianus (Black-tailed Prairie Dog) and 
Neotoma floridana (Eastern Woodrat) but the invasive 
Rattus norvegicus (Brown Rat) has been removed and 
the species of foxes, Martes, and skunks that were each 
previously presented in two chapters have each been 
combined. Furthermore, three chapters have been ex- 
panded including the addition of Macrotus californicus 
(California Leaf-nosed Bat) to the bats, Ammospermo- 
philus (antelope ground squirrel) to the ground squirrels, 
and the subsuming of Cervus nippon (Sika Deer) into 
a more inclusive non-native large mammals category 
covering several species at the end of the book. Recent 
taxonomy also has been incorporated such as the 
generic use of Lontra for the river otter, Puma for the 
Mountain Lion, and Jayassu for the Collared Peccary. 

Of the over 400 species of mammals known from 
North America, approximately half (210) are covered 
but 155 of these species are not full accounts. The 
chapters range from 28 detailed single-species accounts 
to six chapters that focus on two species with multi- 
taxa reports comprising the remainder. Some of these 
latter chapters concentrate on higher taxonomic groups 
including the six species of voles (genus Microtus) 


found in North America; two genera of ground squir- 
rels (Spermophilus and Ammospermophilus) covering 
25 species; six species of foxes in the genera Alopex, 
Urocyon and Vulpes; 19 species of pocket gophers in 
the family Geomyidae; six species of bats from the 
Vespertilionidae family, one species from Molossidae, 
and one species from Phyllostomidae; six species rep- 
resenting the toothed whale suborder Odontoceti; and 
11 species of the baleen whale suborder Mysticeti. The 
final chapter treats several exotic or alien species and 
their associated problems as related to the native fauna. 

The general format within each account essentially 
has remained the same as the first edition. Chapters 
begin by reviewing the nomenclature, distribution, and 
description of the species or species-group. Most 
accounts include life history topics such as physiology, 
reproduction, age estimation, ecology, feeding habits, 
behavior, and mortality. A summary is then presented 
on the economic status, management, conservation, and 
research needs of the taxa under study. Other subjects 
covered by some but not all accounts are genetics, 
anatomy, development and habitat. Except for the last 
chapter, all have distribution maps, skull figures, and 
most have photographs of live animals. The book ends 
with two appendices identifying cranial bones and illus- 
trating standard cranial measurements, a glossary, and 
an index. 

With over 100 contributors to this edited book on 
wild mammal species deemed to have management 
significance in North America, it was inevitable that 
there would be inconsistent treatment across groups. 
For example, the account of the Black Bear (Ursus 
americanus) is one of the shortest chapters although 
it is a relatively well-studied and endangered mammal 
that is in need of a comprehensive management pro- 
gramme. There is almost no mention of its conservation 
status or of its economic importance, and the chapter 
is not much changed from the first edition. In contrast, 
the longest single-species account is for the Bison 
(Bison bison), a highly managed species with very few 
free-ranging individuals. The text has been substantially 
revised from the original account and is one of the 
more thoroughly covered species. Within the multi-taxa 
chapters, the presentation of information was not stan- 
dardized, making it difficult to locate specific infor- 
mation for comparative purposes. For example, the 
seals began with general characteristics for pinnipeds 


2004 


and then finished with individual accounts for the 16 
different species. In contrast, toothed whales included 
three detailed and sequential species accounts fol- 
lowed by abbreviated discussions on three other 
species. 

Overall, this revised edition contains some of the 
most comprehensive descriptions of mammal species 
found in North America. Notwithstanding the afore- 
mentioned criticisms, the editors have continued the 
fine tradition of thoroughness and scholarship estab- 
lished in their first volume. Not only are the accounts 
a summary of our biological knowledge of the larger 
species but the information also is interpreted and 
presented in the context of management and conser- 
vation. This book will be the primary reference source 
that bridges the gap between applied biology and policy 


Book REVIEWS 


473 


implementation. A companion volume dealing with 
the small mammals, and the other half of mammalian 
diversity, in North America would be a nice comple- 
ment but perhaps wishful thinking for others to under- 
take. One noticeable drawback is that the publication 
is expensive, which will discourage students, researchers, 
and government workers from purchasing it for their 
personal library. This book, however, is a required 
monograph for institutional and departmental libraries 
that have an interest in not only mammals and their 
conservation or management but also wildlife ecology, 
in general. 


BURTON K. LIM 
Centre for Biodiversity and Conservation Biology, Royal 


Ontario Museum, 100 Queen’s Park, Toronto, Ontario 
M5S 2C6 Canada 


| Pete Dunne on Bird Watching: The How-to, Where-to and When-to of Birding 


By Peter Dunne. 2003. Houghton Mifflin, 222 Berkeley Street, 
Boston, Massachusetts 02116. USA. 352 pages. U.S. $12. 


Birdwatching is one of the fastest growing outdoor 
activities in North America. Once the obsession of a 
few, it is now firmly lodged in the mainstream. Bird 
books have proliferated, the optical equipment gets 
better every year, and birding clubs and media are now 
commonplace: there are even television shows about 
birding. With all these riches, what is a novice to do? 
Pete Dunne’s latest work is a good place to start. With 
this cleverly thought out book, Pete Dunne starts at 
the beginning and provides the tools and tips to make 
birding a lifelong journey of discovery. 

The author is a gifted communicator and teacher, 
making the material accessible and a fun read, while 
at the same time packing dozens of tools and tips into 
three hundred information-filled pages. The place he 
Starts is the backyard — precisely the place where many 
people first get hooked on birds. He then walks the 
reader through the tools of the trade, the fundamen- 
tals of birding, applied birding (“for fun, purpose... 
| even profit!”), eventually bringing the reader full cir- 
_ cle to ethics and a solid conclusion that reminds us of 
_ why we birdwatch. Each chapter ends with a useful 
_ summary of key learning objectives, which helps to 
_ hammer home the important points. He keeps the 
| material alive by interspersing anecdotes from his own 
rich experience, and others gleaned from a veritable 
who’s who of North American birding. There are plen- 
ty of photos (black-and-white, this is no coffee table 
| book) to illustrate points the author wants to make. 
| The author’s dry wit frequently surfaces, so be pre- 
_ pared for the occasional good laugh. 

Some of the advice is priceless, particularly ten key 
questions to ask when identifying a bird, and a sec- 
tion entitled “learning to see.” This is complemented 
by practical advice on things like how to pick binoc- 
ulars, field guides and spotting scopes. For example, 
_ the author provides a helpful hint on how to check to 


see if that great pair of binoculars you are thinking of 
buying is in alignment; if they are not, your eyes will 
suffer. 

There is also plenty of advice — generic and specific 
— on “where to go” and how to maximize your birding 
once you get there (my favorite: “the power of the 
pause”, wait, and birds will show up). He also talks 
about how to contribute to the store of knowledge 
while having good fun, for example by participating 
on Christmas Counts and Breeding Bird Atlases. In 
the final chapters the author notches the level up sev- 
eral grades, letting novice birders in on some of the 
inner secrets of successful birders like how to be where 
and when the birds are. He even divulges the secret 
of the perfect Eastern Screech Owl imitation (I’m not 
telling, you will have to read the book to find out.) 

Are there things I do not like about this book? Not 
many, but there are a few. The format, with plenty of 
inserts, is occasionally disconcerting, particularly some 
of the “‘insider’s insights” with sometimes abrupt shifts 
from the author’s voice to another, in one case into a 
lecturing tone thankfully absent in the rest of the book. 
Sometimes the order of things is confusing, for exam- 
ple the discussion on spotting scopes is widely sepa- 
rated from that on binoculars. While this follows a 
logical sequence (most birders start with binoculars 
and only “graduate” to scopes later on in their birding 
careers) is does seem misplaced. The advice provided 
is solid and if followed will make for better birders, 
but there are a few minor missteps. For example, the 
author perpetuates the oft-repeated myth that European 
birds are less responsive to squeaking than those in 
North America — not true in my experience. The book 
is also unabashedly North American centric; there is 
very little here about the rest of the world. Occasion- 
ally the book drifts towards the advertorial, for example 
a limited number of bird tour companies are high- 
lighted, but generally the author strikes the right bal- 
ance. There is one point in the book that I objected 


474 


to, and that was the description of the author, in a line 
with a dozen other birders, wading through a long 
grass prairie in an attempt to flush a Baird’s Sparrow. 
While this anecdote was properly set in a discussion of 
ethics (he stepped on a grouse nest) the action described 
was out of character with the rest of the book (or even 
the advice provided in the rest of the book) and the 
ethical dimensions could have been more forcefully 
argued. These points do not seriously detract from the 
book, and I would rank it among the best birding “how- 
to” books I have encountered. 


BOTANY 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


This book is aimed at the novice. If you have just 
started birding, this book is for you. If you know some- 
one who has just started, this book would make a great 
gift. You may even want to lend it to your spouse, 
friends or relatives; anyone who is trying to figure out 
what all the fuss is about. Better still: donate it to the 
local school library after you are done with it. Will 
there be more to learn after you put this book down? 
Absolutely, but that is the whole point of the book: 
birding is a lifelong discovery. 


MARK GAWN 
1354 Viking Drive, Ottawa, Ontario K1V 7J6 Canada 


Alpine Plants of North America: An Encyclopaedia of Mountain Flowers from the Rockies to 


Alaska 


By Graham Nicholls and Rick Lupp, Consulting Editor. 2003. 
Timber Press, 133 SW 2nd Avenue, Ste. 450, Portland, 
Oregon 97204 USA. 344 pages. U.S. $49.95. Cloth. 


Entitled Alpine Plants of North America, this 344 page 
book takes an all-embracing look at many of nature’s 
floral delights, which the author explains as being cate- 
gorized as alpines, though they may be found any- 
where from seaside to high mountain. Mr. Nicholls 
wins my applause by telling us that he likes to limit 
his plants to 30 cm (12 inches) in height, which I find 
keeps plants in scale in an average home rock garden. 

The title might be a bit amusing to a Canadian — it 
appears to have been accomplished by excluding any 
plant references to land we hold dear! We are told the 
book is intended for practical, on site use, both in the 
field and in an owner’s garden — generally Graham 
has succeeded but he falters a bit in the area of plant 
identification — something which is vital to enthusias- 
tic alpine plant lovers. May I suggest that a few less 
species photographs and more emphasis on their clarity 
(there were a number of photos from which identifi- 
cation would be difficult if not impossible). More close- 
ups of flowers and foliage would help a great deal. 

I thought that the format of the book was excellent: 
information on plants’ natural growing areas and cul- 
tural tips made for a most helpful package of useful 
information. Choosing the best location and growing 
medium for our newest acquisitions is very often a 
painful experience. Having several identical plants and 
plenty of spare sites is sometimes our best hope of 


Arboretum America, A Philosophy of the Forest 


By Diana Beresford-Kroeger. 2003. University of Michigan 
Press, 839 Greene Street, Ann Arbor, Michigan 48104- 
3209 USA. 196 pages. U.S. $29.00. 


Arboretum America, A Philosophy of the Forest is 
a unique work. It’s a book about trees in a compound 
context — global, local and personal — informed by an 


success. However, a careful reading of the needs of each 
species covered in this book should save many early 
plant funerals! 

The introduction of little known species is great fun 
and I applaud Mr. Nicholls’s inclusion of Talimums 
in his writing. I have only one species in my collec- 
tion to date — T. selinoides — but after seeing photos 
of such beauties as T. brevifolium, and T: *Zoe’, I shall 
be very soon searching for more. It is most amazing 
to observe the miniature size, quantity and length of 
blooming period in this enjoyable plant. 

It is refreshing to find a proven plantsman willing 
to share his knowledge of plant propagation so freely. 
This is most evident in his detailed directions concern- 
ing the taking of cuttings from various species of Phlox. 
Perhaps his most helpful writings in this area are the 
descriptions of each species’ natural surroundings, soil 
conditions, and moisture tolerance. The book entices 
the reader to find suppliers of seeds, plants, or best of 
all to follow in Graham Nicholls’ footsteps — to see at first 
hand the alpine specimens he has so carefully covered. 

Both author Graham Nicholls and consulting editor 
Rick Lupp have done a magnificent job in producing 
a book of much needed information about the identi- 
fication and growing of alpines and done it in a neat, 
readable, and orderly fashion. This book will give the 
reader true value for his or her money. 


WILLIAM BARKER 


8 Stonecroft Terrace, Kanata, Ottawa, Ontario, K2K 2T9 
Canada 


intriguing variety of perspectives, including ecology, 
ethnobotany, horticulture, ethnology and mythology. 
The result is an eclectic and appealing book, no great 
surprise considering that the author, Canadian Diana 
Beresford-Kroeger, is, according to the back cover, a 
“botanist, medical and agricultural researcher, lecturer 
and self-defined “renegade scientist” in the fields of 


2004 


classical botany, medical biochemistry, organic chem- 
istry and nuclear chemistry.” 

The book starts with an introduction that includes 
appealing stories from Beresford-Kroeger’s child- 
hood, a discussion of the world as a global garden, a 
definition of Beresford-Kroeger’s concept of “bio- 
planning’’(covered in her previous book Bioplanning 
a North Temperate Garden), and a description of how 
bioplanning applies to forests. She then goes on to 
profile 20 North American tree groupings, including 
— genus followed by common name — Acer (Maple), 
_ Asimina triloba (Pawpaw), Betula (Birch), Carya 
(Hickory), Catalpa (Catalpa), Crataegus (Hawthorn), 
Fraxinus (Ash), Gleditsia (Honey Locus), Juglans 
nigra (Black Walnut), Magnolia acuminata (Cucum- 
ber Tree), Ostrya virginiana (Hop Hornbeam), Pinus 
(Pine), Ptelea trifoliata (Wafer Ash), Quercus (Oak), 
_ Sambucus (Elderberry), Sassafras (Sassafras), Thuja 
occidentalis (Cedar), Tilia (Basswood), Tsuga (Hem- 
| lock), Ulmus (Elm). 

It’s puzzling, this choice of trees. Why these group- 
ings and not others? Why include elderberries and 
leave out dogwoods, why hemlocks and not firs, why 
_ birches and not aspens? The author never tells us why; 
_ never explains her inclusion and exclusion criteria. 
And I can’t help but wonder, as I wander the fields 
_ and forests of home, about the cherries and beeches, 
_ tamaracks and spruces ... should they not be includ- 
ed in forest bioplanning too? 

That little grievance aside, the book is vibrant and 
_ delightful, with much to teach from the wide range of 
_ perspectives mentioned earlier. Each profile includes 
_ six sections: “The Global Garden”, tracing the history 
and geography of each tree grouping; “Organic Care”, 
_ covering related horticultural topics; “Medicine”, 
discussing traditional and potential medicinal uses; 
“Ecofunction”, describing the trees’ ecological roles; 


_ Cape Cod Wildflowers: A Vanishing Heritage 


By Mario J. DiGregorio and Jeff Wallner. 2003. University 
Press of New England, One Court Street, Lebanon, New 
Hampshire 03766 USA. 169 pages. U.S. $19.95. 


_ This is a reprint of the original volume first pub- 
_ lished in 1989 by Mountain Press Publishing Company, 
_ with a new introduction by the authors. As the authors 
| state, “This book is an appreciation, a guide, and a 
_ plea for protection. On one hand it explores the arcane 
_ lore of the medieval herbalists... On the other, it chron- 
icles the latest scientific understanding of flowers’ 
ecological importance and current attempts to pre- 
serve natural diversity.” 

If not unique, this book is at least one of very few 
examples of this particular approach to botanical 
description. It is arranged in chapters based upon habitat 
types, all common to the Cape Cod area. Each habitat 
is described in the opening of its chapter. The habitats 
_ include woodlands; ponds and bogs; sandplains; salt 


Book REVIEWS 


475 


“Bioplan”, explaining how the trees have been and 
could continue to be incorporated into human envi- 
ronments; and “Design”, about the trees’ appearances 
and aesthetic properties. 

Much of the information was new to me. Like a 
maple biochemical, acerin, currently being investi- 
gated for antiviral and antibiotic qualities. Like the 
birch being one of the two sacred trees given to the 
Aboriginal peoples of North America. Like pines con- 
tributing numerous important air-freshening com- 
pounds to the atmosphere. Like hawthorn fruit being 
called a “pome” which, along with the leaves, pro- 
duces a number of biochemicals which form a high- 
energy compound beneficial to migrating birds. Like 
basswoods producing huge crops of nectar at a time 
when bees need it desperately and many other flow- 
ering plants have stopped blooming. 

The profiles are interspersed with photographs of 
other members of the forest community — lichens, 
fungi, flowering plants, shrubs — which help enhance 
the bioplanning concept and paint a larger picture of 
the forest ecosystem. And the occasional yellow boxes 
with relevant stories from the author’s life add a pleas- 
ing personal touch. 

One of the most personal touches in the book appears 
at the end of the introduction, where Beresford-Kroeger 
shares with us her dream “that a moratorium will be 
put on the cutting of what is left of the global forests 
and that ordinary people with an acorn and a shovel 
begin the long road back to nature.” Ordinary people. 
That’s me. Though I don’t know if I have any acorns 
around. But there’s a collection of shovels in the gar- 
den shed, and space along the fence. Perhaps I could 
manage to find a handful of maple keys ... 


R. SANDER-REGIER 
RR5 Shawville, Quebec JOX 2YO Canada 


marshes; and dunes and beaches, with an additional 
chapter to accommodate alien species. 

Each habitat type is represented by ten to fifteen 
species endemic to each area. As the authors admit, 
the sampling is limited in scope, covering only 66 of 
the 1300 species of vascular plants found in the Cape 
Cod area. Each species is covered on facing pages, 
with the left-hand page containing a description of the 
plant and other comments on such aspects as propa- 
gation and threats to its environment. A colour photo- 
graph of the flower is located on the right hand page. 

The authors could have, however, spent a little more 
time in researching their data, especially since this is 
the second edition of their work. The pollination process 
of Cypripedium acaule, for instance, suffers from the 
following extremely fanciful and highly inaccurate 
description. “Drawn by the promise of a sugar “high” 
from the nectar inside, insects enter through the slit... 
On the way out the insect is plastered with a natural 


476 


glue on which, in turn, pollen is dusted. Emerging from 
the flower with cargo (of pollen) and payment (in nec- 
tar) the insect goes to another flower where, in the same 
intricate exit process, some of the pollen is deposited 
on the stigma to fertilize the plant and produce new 
seed.” Of course, Cyp. acaule has no nectary, therefore 
there is no payment in nectar; pollen is not dusted; 
and the picture of an insect “plastered with natural 
glue” is entirely missleading. Because the above pas- 
sage occurs in chapter one, a suspicion tends to be 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


planted early as to the accuracy of the rest of the text. 

The authors are to be commended in producing a 
book dedicated to the ever growing threats to our nat- 
ural environment. Although the scope of this volume 
is limited to the Cape Cod area of Massachusetts, its 
message could equally apply to almost anywhere in 
North America. 


WILLIAM R. ARTHURS 


1228 Lampman Crescent, Ottawa, Ontario K2C 1P8 Canada 


Lewis Clark’s Field Guide to Wild Flowers of the Sea Coast in the Pacific Northwest 


Compiled and photographed by Lewis J. Clarke, edited and 
composed by John G. Trelawny. 2004. Harbour Publishing 
P.O. Box 219, Madeira Park, British Columbia VON 2HO 
Canada. 80 pages. Second Edition.Canadian $12.95. 


The first edition of this beautiful little book was pub- 
lished by Gray’s Publishing Limited, Sidney, British 
Columbia, in 1974 and numbered 64 pages. In this sec- 
ond edition which numbers 80 pages, John Trelawny 
has made some changes in the introduction and includ- 
ed some acknowledgements. Like the first edition, 
the pages are not numbered, but each species and the 
accompanying descriptive text is numbered sequen- 
tially so that they can readily be found from the index 
which includes both scientific and common names. 

The 95 flower pictures in this second edition are 
absolutely beautiful and in many are somewhat clearer 
than those in the first edition because of more modern 


technology in producing them from the photo slides 
and like the 1974 edition each photo has a scale mark- 
er which gives the size of the picture in relation to the 
average plant size. A glossary at the end of the volume 
depicts various leaf shapes and flower parts to aid the 
user in understanding the descriptive text which ac- 
companies each photograph. New plant photographs 
have been provided by T. & S. Armstrong, M. Barker, 
Ugo Cagnetta, W. Merilees, F. Pratt, H. Roemer, W. 
van Dieren, and M. Wheatley. 


WILLIAM J. Copy 


Biodiversity, National Program on Environmental Health, 
Agriculture and Agri-Food Canada, Research Branch, Wm. 
Saunders Building, Central Experimental Farm, Ottawa, 
Ontario K1A 0C6 Canada. 


Wild Flowers of Field & Slope in the Pacific Northwest 


By Lewis J. Clark. 2002. Harbour Publishing, P.O. Box 219, 
Madeira Park, British Columbia VON 2HO Canada. 108 
pages.Canadian $9.95. Paper 


Wild Flowers of Forest & Woodland in the Pacific Northwest 


By Lewis J. Clark. 2003. Harbour Publishing, P.O. Box 219, 
Madeira Park, British Columbia VON 2HO Canada. 100 
pages. Canadian $12.95. 


Wild Flowers of the Mountains in the Pacific Northwest 


By Lewis J. Clark. 2003. Harbour Publishing, P.O. Box 219, 
Madeira Park, British Columbia VON 2HO Canada. 106 
pages. Canadian $12.95. 


These three books contain absolutely beautiful colour 
photographs. The photographs are numbered sequen- 
tially and each one has a marker [0.5] to indicate its 
size and is accompanied by a printed paragraph with 
the common and scientific names, a detailed descrip- 
tion, together with the habitat and range, and a number 
to indicate its sequence in the book. There are 108 in 
the first book, 100 in the second and 106 in the third. 
There is a four or five page interesting introduction at 


the front of each book and an index, glossary and a 
list of Additional Field Guides available from Har- 
bour Publishing at the end. There are most interest- 
ing pictures on the front covers of each book. On the 
back covers there is a note about the author, Dr. Lewis 
J. Clark together with a map of the Pacific Northwest 
on which there are shaded areas depicting where the 
wild flowers can be found. All are elegant. 


WILLIAM J. Coby 


Biodiversity, National Program on Environmental Health, Agri- 
culture and Agri-Food Canada, Wm. Saunders Building, Cen- 
tral Experimental Farm, Ottawa, Ontario K1A 0C6 Canada 


2004 


ENVIRONMENT 


BooK REVIEWS 


477 


Conserving Living Natural Resources in the Context of a Changing World 


By Bertie Josephson Weddell. 2002. Cambridge University 
Press, 40 West 20th Street, New York, New York 1001 1- 
4221 USA. 426 pages. £70 Cloth, £24.95 Paper. 


Conserving Living Natural Resources in the context 
of a changing world is an excellent information source 
for anyone interested in natural resource management; 
in the evolution of attitudes, mainly North American, 
and knowledge regarding natural resources; in the 
development, over time, of the relationship between 
human beings and the natural environment — or any 
combination of the above. 

Weddell writes in the preface that she designed the 
book as an introduction to natural resource conserva- 
_ tion for students, and as a review for managers. As such 
' it covers three different approaches to natural resource 
_ management — utilitarian, preservationist, and sustain- 
able-ecosystem — in more or less chronological order. 

For each approach, Weddell traces the historical 
conditions that set the stage for that type of resource 
management and discusses its strengths and weakness- 
es. She also explains the approach’s central concepts, 
both philosophical and scientific, and describes its 
principal techniques. Weddell makes sure to empha- 
size why it is important to learn about the different 
approaches, even if certain aspects are now slightly 
outmoded. She makes four excellent points: (1) that 
there is no single correct way to manage natural re- 
sources; (2) that our generation has not necessarily 
discovered “the truth”; (3) that it is important to under- 
stand how we got to where we are today so that we 
can learn from our mistakes; and (4) that some of the 
approaches discussed in the book, even if they are 
out of date, are still widely applied today. 

Part I of the book, “Management to maximize pro- 
duction of featured species — a utilitarian approach to 
conservation’, discusses the commodification of re- 
sources, the impacts (habitat alteration, species de- 
clines, extinctions) of the commodification, and the 
responses (regulation, protection) to those impacts. It 
_also traces the development of the discipline of natu- 
ral resource management, describes the central concepts 
of the utilitarian approach (population growth, inter- 
actions between populations, habitats), and explains 
_ its main techniques (harvest management, habitat man- 
agement, management to minimize conflicts between 
pest species and people). 

In Part I, “Protection and restoration of populations 
_ and habitats — a preservationist approach to conserva- 

tion”, Weddell describes economic and demographic 


changes after World War II, and discusses the in- 
creasing awareness of ecological problems during that 
period (invasive species, toxic substances, ongoing 
extinctions). She also covers the rise of preservationist 
management, its central concepts (causes of extinction, 
speciation, classification), and the principal techniques 
relating to species protection and restoration, and to 
ecosystem protection and restoration. 

In Part ILl, “Management to maintain processes 
and structures — a sustainable-ecosystem approach to 
conservation”, Weddell provides an overview of more 
recent pressures to move beyond the protection of 
species and reserves, including practical, scientific, 
political, ethical and philosophical considerations. She 
also traces the rise of sustainable-ecosystem manage- 
ment, describes its central concepts (equilibrium the- 
ories, the flux of nature), and discusses two main tech- 
niques: conserving natural processes and contexts, and 
including people in the conservation process. 

The book covers a lot of ground, much of it fairly 
complex, and it isn’t short. But Weddell’s clear and 
simple writing style makes the content is easy to read 
and understand. And she includes lots of helpful dia- 
grams and dynamic examples. It all makes for -pleas- 
ant and highly informative reading — particularly the 
historical background sections, which provide a valu- 
able and relevant context for understanding the disci- 
pline of natural resource management. 

Weddell closes the book by emphasizing that each 
of the natural resource management approaches — utili- 
tarian, preservationist and sustainable-ecosystem — has 
much to teach us about solving contemporary problems. 
She stresses that each approach has advantages and 
disadvantages that make it appropriate to particular 
circumstances, and that it is possible to blend elements 
from all three approaches. 

She also points out, and I quote: “I believe that as 
we continue to search for responsible ways to manage 
living natural resources, a large dose of humility is 
appropriate. Science, whether theoretic or applied, is 
an ongoing process.... Although our understanding of 
the natural world is more detailed than it used to be, 
there is still a lot we do not know. Management should 
err on the side of caution, therefore. There will always 
be surprises.” Wise words. 


R. SANDER-REGIER 


RRS5 Shawville, Quebec JOX 2YO Canada 


478 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Visions of the Land: Science, Literature, and the American Environment from the Era of 


Exploration to the Age of Ecology 


By Michael A. Bryson. 2002. The University of Virginia Press, 
Box 400318, Charlottesville, Virginia 22904-4318 USA. 
228 pages. U.S. $16.50. Paper. 


Visions of the Land, part of an “Explorations in 
Ecocriticism” series, is an interesting look at the rela- 
tionship between science, the natural environment and 
human beings, as expressed in literature published in 
the United States from, as the subtitle suggests, the 
period of exploration in the 19" century to roughly 
the 1960s. Some aspects of that relationship would 
also apply to Canada. 

The authors examined in the book cover quite the 
range — from explorers John Charles Frémont (Amer- 
ican West) and Richard Byrd (polar regions), to author 
and feminist Charlotte Perkins Gilman, naturalist Susan 
Fenimore Cooper, and scientists John Wesley Powell, 
Rachel Carson and Loren Eiseley. The types of writ- 
ing also vary widely — from exploration narratives 
and technical reports, to fictional utopias, natural his- 
tories, popular scientific literature, and more. 

The book is divided into three parts: |. Narratives 
of Exploration and the Scientist-Hero (Frémond and 
Byrd); 2. Imagined Communities and the Scientific 
Management of Nature (Powell and Perkins Gilman); 
and 3. Nature’s Identity and the Critique of Science 
(Fenimore Cooper, Carson and Eiseley). The book’s 
sub-title suggests a certain chronological order in the 
material, but that turned out not to be the case, which 
made reading and comprehension a little confusing. 
It was also a little difficult to follow the author’s argu- 
ments through the themes covered in the three parts. I 
would have found it more effective had Bryson sim- 
ply written a chapter on each of the authors. 

Nevertheless, there is lots of good content in the 
book regarding science and connections with human 
perspectives on the natural environment. Frémont, for 
example, was the “glamour boy of American westward 
expansion”, the archetypal macho scientist-explorer, 
who worked and wrote both scientific reports and 
popular literature at a time when the country was ex- 
panding ever faster westward, and when surveying 
technology and cartographic techniques were contin- 
ually improving. Frémont’s writing combines descrip- 
tive and poetic elements with analytical, quantitative 


elements. The science he practises and expresses is 
highly rational and empirical, and the land something 
to be studied, catalogued and mapped. 

John Wesley Powell, featured in Part 2 of the book, 
is another fascinating character in the history of the 
American West. An explorer-scientist like Frémont — 
as well as an ethnologist, philosopher, writer and gov- 
ernment leader — Powell was, according to Bryson, 
“one of the most important and influential scientists 
of his age.” Powell’s work emphasises the scientific 
control of a mechanistic nature while at the same time 
recognising nature’s self-regulating properties and the 
need for careful settlement and agricultural practices. 
Powell is aware of and interested in community, par- 
ticularly with respect to the need for human commu- 
nities to develop a responsible relationship with nature 
and to use science wisely. 

Susan Fenimore Cooper, featured in Part 3, is one 
of the two non-scientist writers discussed by Bryson, 
although she was a dedicated naturalist and keen 
observer. Her work falls into the 19" century natural 
history writing tradition which helped inform the 
ecological science developed in the following century. 
Her book, Rural Hours, published in 1854, combines 
natural history, cultural analysis and personal stories 
to create an environmental and social portrait of her 
home region in central New York state. Bryson finds 
Rural Hours fascinating “not only because it com- 
bines multiple strands of the nature writing tradition 
but also because it provides a complex and fairly sub- 
stantial critique of the relation between nature and 
the human community.” Fenimore Cooper views sci- 
ence not as something which can be used to objectify 
or control nature, unlike Frémont or Powell. Science, 
in her view, is a “system of study meant to foster 
moral and intellectual connections between the 
observer and the outside world.” 

I learned a lot about the evolution of scientific 
thinking and its influence on the human-nature rela- 
tionship in Visions of the Land. 1 also discovered some 
fascinating writers and books — ones I’m sure to look 


up and enjoy in the future. 
R. SANDER-REGIER 


RRS5 Shawville, Quebec JOX 2Y0O Canada 


Genetics, Demography and Viability of Fragmented Populations 


Edited by A. G. Young, and G. M. Clarke. 2000. Cambridge 
University Press, 40 West 20th Street New York, New York 
10011-4221 USA. 438 pages. Canadian $63. Paper. 


This is another Australian CSIRO (Commonwealth 
Scientific and Industrial Research Organization) flag- 
ship publication of international importance to research 
and conservation. “The continuing global trend towards 
non-sustainable exploitation of natural resources means 
that more, rather than fewer, species are going to be 
affected by habitat loss, degradation and fragmenta- 


tion in the future”. The editors of this important book 
make it clear that the species of this globe will either 
have to survive in zoos, or in smaller fragments. For in- 
stance, the range of many Australian mammals declined 
already by over 90% and is now confined to off-shore 
islands; and in New Zealand the native forest cover has 
already been reduced from 78% in pre-human times to 
23% today. Obviously, for many plants and animals 
in the world, preservation with relatively intact habitats 
is no longer an option. 


2004 


Already Darwin (1876) had documentated inbreed- 
ing depression in both cultivated and native plant spe- 
cies. The two major goals of modern conservation gen- 
etics are the maintenance of genetic variation and the 
avoidance of inbreeding depression. 

This book consists of three parts (Introductory Con- 
cepts, Animal and Plant Case Studies) and includes 
20 research papers from 42 international authors. Most 
of the contributors come from the southern hemi- 
sphere (Australia and New Zealand) as well as from 
the English speaking research sphere (USA, UK). 
However, their research applications come from many 
parts of the world and cover the variety of the animal 
kingdom (Red-cockaded Woodpecker, Golden Lion 
Tamarins, Mexican Wolves, Bonytail Chub, Chinook 

~ Salmon, Golden Sun Moth, Desert Bighorn Sheep, 
small mammals) and a variety of plant species. The 
plant section is of special interest here as plants are 
used to present genetic mechanisms transferable to 
animal populations and their conservation. It even 
includes an endangered dry forest tree species (Swi- 
etenia humilis) in Central America now listed with 
CITES (as are two Mahagoni tree species; formerly of 
major economical value). I also admire the 54 pages 
_ of references and a detailed book index. 

Although local extinction is not infrequent to observe 
in nature “Habitat destruction is the most obvious cause 
of species decline and the most difficult to reverse”. 
Recent fragmentation shaped the global landscape 
nowadays, which has nothing to do anymore with the 
original habitat. This books makes a pledge that we 
need to find a limit how we affect the global environ- 
ment. “The techniques for doing so will come more 
from economics, psychology and sociology rather than 
from biology”. However, as the editors show, biology 
has an important role to play beyond identification of 
species and ecosystems. 

It’s a challenge to find the right gene that determines 
inbreeding in a cost-effective way. The book is full of 
GGTGCTAGs, Dendrograms, Nei Genetic Distances, 
Phylogenies etc. The presented papers report in great 
detail on DNA extraction and processing methods such 
_as PCR, but less so on statistics. Commonly applied 
software packages and methods to derive evidences are 
|GENEPOP, GenAIEx, SPARKS, PROC INBREED 
(SAS), PCA, bootstrapping, VORTEX and RAMAS. 
Authors conclude that relying entirely on neural molec- 
ular markers is inadequate. Most of the presented gene 
studies in this book are based on mtDNa and Allo- 
zymes; microsatellites might track fragmentation pro- 
cesses quicker. 

_ Concepts and terms like Minimum Viable Popula- 
tions (MVP) metapoluation paradigm, effective pop- 
ulation size, rescue effect, simulation models, demog- 
-taphy, transition matrix and the movements of pests, 
pathogens and predators are discussed as well. Some 
of the presented studies cover a period of over 10 years. 

The well known study on Cheetah inbreeding effects 
(e.g., population decline, susceptibility to disease) 
‘Temains controversial. But in Bighorn Sheep, high 


Book REVIEWS 


479 


heterozygosity is known to be associated with large 
horn size at sexual maturity, which confers breeding 
superiority. It is worthwhile to note that over the last 
200 years alone 98% of Desert Bighorn Sheep popu- 
lations have been reduced. 

This complex publication makes for a bible on 
fragmentation as being genetically harmful. 

Much of the book deals with the landmark publi- 
cations by Caughley (1994) and Lande (1988) and pro- 
vides further evidence and research on these topics. It 
is in support of the classical conservation arguments; 
e.g., Those brought forward by Soulé. The book some- 
what promotes the Australian conservation view, which 
might not necessarily be shared world-wide. For in- 
stance, the foreword cites Randal O’ Toole (1999) who 
stated that “conservation biology is not a science but 
a political movement based at least in part of nine- 
teenth-century ideals of what an ecosystem is all 
about”. The reader of this book will definitely learn 
that genetic loss will have negative effects on econom- 
ic and physical well-being. “Each extinction erodes 
the biological legacy, and humanity slips away. Stew- 
ardship for the planet and its inhabitants will have 
been lost, replaced by greed, ignorance, and short- 
sightedness”. Brooks (1997) contemplated already that 
“while the ship is sinking, conservation geneticists are 
busy counting the deck chairs”. The contributing authors 
to this book make a case for managing through genet- 
ics; they are frank about the fact that many problems 
with endangered species cannot be solved with 
genetic techniques alone but that genetics will allow 
for much more informed decisions. As the authors 
show, true population estimates are hard to get from 
field work, and DNA offers major opportunities to infer 
some basic life-history, demographic and population 
parameters. 

I am very glad that this book stresses the impor- 
tance of spatial issues for PVA, but true GIS papers 
are unfortunately not presented. Some papers deal 
indirectly with what can be defined as Landscape 
Ecology though. Readers will also appreciate the great 
summary of the most important elements of a suc- 
cessful PVA. Some really good model philosophy is 
presented, too. PVAs are coined as a “loaded gun” 
pointing towards economics and others. The book 
deals nicely with data uncertainties in PVAs, PVA 
acceptance in the research community and PVA appli- 
cations, too. “Incomplete information does not mean 
that meaningful results are impossible to obtain because 
there is very significant value in building a model for its 
own sake”, e.g., as a guide for further data collection 
and bringing stakeholders together. A key argument 
is that a PVA compares risk, but does not measure it. 

Besides the many other interesting and important 
papers in this book I, personally, really like the one 
dealing with the faunal collapse and genetic erosion 
in small mammal communities on islands in the Chiew 
Larn reservoir in Thailand: it’s an island habitat mile 
stone study with a 10 year data set. 


480 


The presented plant studies are very valuable because 
plants as well as their response to habitat loss and frag- 
mentation, are usually understudied. The pollination 
papers show nicely that fragmentation has to consider 
pollinators (bees and butterflies), too, besides the pollen- 
mediated gene flow. 

The correlation between population size and genet- 
ic diversity is clear. This book supports the view that 
it is increasingly apparent that the consequences of 
fragmentation are dependent upon complex interactions 
of demographic and genetic variables. However, the 
relative importance of demography vs. genetics for 
conservation of small populations or rare species is 
still debated. In either case, inbreeding and reproduc- 
tion failure alone have limited demographic effects. 
However, when both are combined, and certainly when 
applying the “precautionary principle” in conservation, 
the two result in significantly reduced population 
persistent times. The authors show well that multidis- 
ciplinary approaches are crucial but that there is a 
large degree of discipline rivalry perpetuated by the 
structure of university faculties and departments and 
by the patterns of research funding worldwide. DNA 


A Primer of Ecological Genetics 


By Jeffrey K. Conner and Daniel L. Hartl. Sinauer Associates, 
Inc., Publishers, 23 Plumtree Road P.O. Box 407, Sun- 
derland, Massachusetts 01375-0407 USA. 304 pages. 
U.S. $47.92. 


Recent developments in molecular biology have 
lead to the application of increasingly sophisticated 
genetic techniques to ecological questions. A primer 
of ecological genetics serves as an introduction to 
this rapidly expanding field. The intended readers for 
this book are advanced undergraduate and graduate 
students who will find it a valuable resource in their 
studies. The authors also suggest that professional 
biologists will benefit from the material presented, 
but I don’t think this audience is as well served. 

The book starts with the basic concepts of population 
genetics, including a review of the most commonly 
used molecular markers. Conner and Hartl then guide 
the reader through theoretical and empirical advances 
gained from two complementary approaches to this 
field. The first is the study of genetic variation in nat- 
ural populations. Observed levels of genetic variation 
are routinely used to assess breeding systems, migra- 
tion, and differentiation within and among popula- 
tions, making a sound understanding of this material 
essential for anyone working in conservation biology. 
The second approach, quantitative genetics, deals with 
insights gained from experimental studies. While much 
of the relevant literature addresses breeding programs, 
the authors argue convincingly for the value of quan- 
titative genetics in studying natural populations. 

A detailed discussion of natural selection follows, 
and the closing chapter of the book covers some of 
the major applications of population genetics — con- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


studies must remain incomplete as long as prey, pre- 
dation, landscape and other factors get ignored. 
Unfortunately, data reported upon in this book are 
not freely available in a digital format for the public, 
nor is that fact anywhere in this book emphasized; the 
important topic of Metadata is not mentioned either. 
This does not add to transparent decision making in 
conservation and should be considered in any study. 
As this book once more shows very clearly, “cur- 
rently, we do have tools available to us that we are not 
using”. If there is something to criticize in this book 
then: Why do we produce, and focus on, superb sci- 
ence, whereas most of our governments and the public 
are over 15 years behind in their actions and implica- 
tions towards efficient conservation? In the meantime, 
the global landscape will be further exploited, with book 
shelves getting bigger. This excellent book calls for 
action: Management considering genetics is required. 


FALK HUETTMANN 


Institute of Arctic Biology, Biology & Wildlife Deptartment 
Fairbanks, University of Alaska, Fairbanks, Alaska 99775 
USA. 


servation genetics, evolution of invasive species, trans- 
gene escape, and the evolution of pesticide resistance. 
This last chapter is an excellent grounding for the 
book, providing concrete examples of the value of 
ecological genetics to real world problems. 

Overall the book is well thought out, and the authors 
succeed for the most part in presenting this complex — 
subject in an accessible way. The prominence of quan- — 
titative genetics in the text is refreshing. The demands 
required for quantitative genetic study have limited its 
use in a conservation context. Conner and Hartl may 
persuade some researchers to rise to the challenge and 
combine observational and experimental work. The 
determined individuals that do will likely produce some 
very interesting results! 

Some of the discussion was over-simplified. The 
reader gains a general understanding of the concepts, 
but will need to refer to the primary literature or a more 
detailed reference to truly master the material. This is 
the authors intention, as evidenced by the excellent 
selection of current references included with each 
chapter, accompanied by discussion questions. In this 
light, the book will make a marvellous study guide for 
a graduate seminar or senior undergraduate course. 
However, professional biologists and conservation 
managers are less likely to have the time and energy 
to devote to the subject. Expanding the treatment to 
make this book a “one-stop” reference would better © 
serve this audience. 


TYLER SMITH 


155 Vanguard Avenue, Pointe Claire, Quebec H9R 3T4 Canada 


2004 


MISCELLANEOUS 


Book REVIEWS 


48] 


Smithsonian Institution Secretary Charles Doolittle Walcott 


By Ellis L. Yochelson. 2001. Kent State University Press, 307 
Lowry Hall, P.O. Box 5190, Kent, Ohio 44242 USA. 590 
pages. U.S. $49.00. 


This, the second and final volume of Yochelson’s 
massive “insider” biography (Yochelson retired from 
the Smithsonian in 1985), is clearly a labour of love. 
The first volume, Charles Doolittle Walcott, Paleontol- 
ogist, appeared in 1998. From Walcott’s daily diaries, 
an unusual amount of detail is provided chronologi- 
cally, month by month, as a record for future historians. 
Fifty pages describe happenings in 1927 alone, when 
Walcott was already seventy years old. 

Charles Doolitte Walcott was a man of humble and 
inauspicious beginnings, without any opportunity for 
a college education. In his early twenties he sold a 
collection of fossils to Louis Agassiz at Harvard Uni- 
versity; a few years later he was offered a job as a 
temporary field assistant, measuring the thickness of 
rocks in Bryce Canyon, Utah, for the newly formed 
United States Geological Survey. A man of great tact 
and wisdom, Walcott never looked back; in 1933 the 
fifth edition of American Men of Science listed him 
posthumously as the third most important geological 
scientist in the country, and he collected twelve hon- 
orary degrees from as many universities. 

Walcott is of enormous importance to Canada. He 
spent about three months each summer for 18 of 19 
years, 1907 through 1925, studying geology and pale- 
ontology in the Canadian Rocky Mountains. In Yoho 
National Park, above Emerald Lake, he discovered 
the Burgess Shale, rated world-wide by Stephen Jay 
Gould as “the most precious and important of all fos- 
sil localities.” In addition to his heavy administrative 
duties during the remainder of the year, Walcott spent 
his spare hours in winter studying and describing 
these fossils. This lifelong devotion to his field of sci- 
ence is all the more remarkable when one considers 
his burdensome official duties. From 1907 until his 
death in 1927 he occupied the powerful post of secre- 


tary of the Smithsonian Institution. He also served 


variously as president of the Paleontological Society, 
National Parks Association, American Philosophical 
Society, Washington Academy of Sciences, National 
Academy of Sciences, and the American Association 
for the Advancement of Science, was chair of both the 
Carnegie Institute of Washington and the National 
Advisory Committee for Aeronautics, and was First 
Vice Chairman of the National Research Council and 
the American Federation of Arts. One of the most 
powerful men in Washington, a mover and a shaker, 
Walcott dined with presidents, cabinet members, 
supreme court justices, and foreign diplomats. In spite 
of these pressures, Walcott was a devoted husband 


and father. His chauffeur, butler, furnace stoker, and 
loyal family retainer at home in Washington, Arthur 
Brown, was also his camp manager, nurse, cook, 
general guardian and friend in each of his mountain 
camps, and the “unofficial mayor of black Washington.” 

Walcott’s first wife, Lura, died of tuberculosis less 
than two years after their marriage. Thirteen years 
later, in 1888, he married Helena Stevens; their hon- 
eymoon was spent looking at rocks in Newfoundland. 
Helena was a competent geologist who, at her first 
International Congress of Geology in London, Eng- 
land, corrected the reversal of Middle and Lower 
Cambrian levels previously described for New York 
State rocks. According to daughter Helen’s reminis- 
cences, her mother Helena was also the first to split 
open a piece of Burgess shale, lying by the trail, while 
waiting for her husband to climb the scree above. It 
proved to have an unusual diversity and complexity 
of fossils, included many soft-bellied organisms, 
unequalled anywhere in the world. Helena had four 
children, all but the oldest of whom accompanied 
them on one or more summer geology expeditions to 
Canada. Helena died in a train accident in Connecticut 
in July 1911, so Walcott had a much shortened field 
season that year. 

In 1914, Walcott married Mary Vaux, shortly before 
her fifty-fifth birthday. A Philadelphia Quaker, she had 
measured and photographed, with her father, George 
Vaux, the movements of Illecillewaet and other gla- 
ciers in the Canadian Rockies, since the 1880s. Mary 
took the place of Helena in assisting with fossil col- 
lecting, but also painted superb portraits of wildflowers. 
Towards the end of his life, Walcott assisted Mary 
Vaux Walcott in having the Smithsonian Institution 
publish her sumptuous five-volume North American 
Wild Flowers, “the monumental ... Audubon of the 
floral world.” The alpine flowers were painted while 
with her husband in the Canadian Rockies. 

Yochelson inserts chatty comments about many top- 
ics, including Walcott’s children. Citations are satis- 
factory, but index entries are haphazardly incomplete. 
I detected nine spelling or typographical errors. 

In the final chapter, Yochelson succinctly and con- 
vincingly counters some of the grossly unfair impres- 
sions left after reading Stephen Jay Gould’s best-selling 
Wonderful Life: The Burgess Shale and the Nature of 
History. Contrary to Gould’s intimations, Walcott sup- 
ported the concept of evolution, “an act of considerable 
courage, not an indication that he was hidebound in 
his thinking ... Walcott did not propose new classes 
and new phyla. The time was simply not ripe ... 
Gould has judged Walcott by the context of his own 
notions more than half a century later.” Walcott’s 


detailed descriptions of the fossil organisms in the 
Burgess shale clearly ranks as among the greatest 
discoveries in paleontology. 

Any reader of this review who has read Gould’s 
Wonderful Life, should read Yochelson’s final chapter, 
and each library that holds a copy of Gould,should 
add this book as well as Simon Conway Norris’ The 


NEw TITLES 
+Available * Assigned 


Zoology 


Amphibians and Reptiles of Pakistan. By M. Khan. 2005. 
Krieger Publishing, P.O. Box 9542, Melbourne, Florida 32902 
USA. No price available. 


The Amphibians and Reptiles of El Salvador. By G. Kohler, 
M. Vessely and E. Greebaum. Krieger Publishing, P.O. Box 
9542, Melbourne, Florida 32902 USA. No price available. 


The Anatomy of Reptiles. By J. Wyneken. 2005. Krieger 
Publishing, P.O. Box 9542, Melbourne, Florida 32902 USA. 
No price available. 


*Birds of Australia —- Seventh Edition. By K. Simpson and 
N. Day. 2004. Princeton University Press, 41 William Street, 
Princeton, New Jersey, 08540-5237 USA. 392 pages. U.S. 
$39.50. 


Birds of the Raincoast. H. Thommasen and K. Hutchings. 
2004. Harbour Publishing, P.O. Box 219, Madeira Park, 
British Columbia VON 2HO Canada. 200 pages. Can $44.95. 
Cloth. 


*A Citizen’s Guide to Ecology. By L. Slobodkin. 2004. Oxford 
University Press Canada, 70 Wynford Drive, Don Mills, On- 
tario, M3C 1J9. 245 pages. U.S. $14.95. Paper. 


Dictionary of Herpetology. By H. Lillywhite. Krieger Pub- 
lishing, P.O. Box 9542, Melbourne, Florida 32902 USA. No 
price available. 


The Exotic Amphibians and Reptiles of Florida. By W. 
Meshaka, B. Butterfield and B. Hague. 2004. Krieger Publish- 
ing, P.O. Box 9542, Melbourne, Florida 32902 USA.166 pages. 
U.S. $34.50. 


Experimental Approaches to Conservation Biology. Edit- 
ed by M. S. Gordon and S. M. Bartol. 2004. University of 
California Press, 2120 Berkeley Way, Berkeley, California 
94704-1012 USA. 358 pages. U.S. $75.00. 


A Field Guide to the Reptiles and Amphibians of Bali. By 
J. L. McKay. 2005. Krieger Publishing P.O. Box 9542, Mel- 
bourne, Florida 32902 USA. No price available.Guide to 
the Amphibians and Reptiles of Japan. By R. Goris and N. 
Maeda. 2004. Krieger Publishing, P.O. Box 9542, Melbourne, 
Florida 32902 USA. 296 pages. U.S. $69.50. Cloth. 


*Mammals of the World — A Checklist. By A. Duff and A. 
Lawson. 2004. Yale University Press, P.O. Box 209040 New 
Haven, Connecticutt 06520-9040 USA. 312 pages. U.S. $45.00. 


+The Monarch Butterfly — Biology and Conservation. By 
K. Oberhauser and M. Solensky (Editors). 2004. Cornell 
University Press, 512 East State Street, Ithaca, New York 
14850 USA. vii + 248 pages. U.S. $39.95. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Crucible of Creation as a rebuttal. On its own merits, 
Yochelson’s book deserves to be in every major library. 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N OJ8 
Canada 


Our Life with Birds. By J. and G. Tveten. 2004. Texas A&M 
University Press, Lewis Street, 4354 TAMU, College Station, 
Texas 77843 USA. 256 pages. U.S. $24.95. Cloth. 


The Panther Chameleon. By G. Ferguson, J. Murphy, J. 
Ramanamanjato and A. Rasselimanana. 2004. Krieger Publish- 
ing P.O. Box 9542, Melbourne, Florida 32902 USA. 166 pages. 
U.S. $39.50 Cloth. 


Penguins of the World. By W. Lynch. 2004. Firefly Books 
Ltd., 3680 Victoria Park Avenue, Toronto, Ontario M2H 3K1 
Canada. 144 pages. Can $35.00. Cloth. 


+ Prairie Ghost — Pronghorn and Human Interaction in 
Early America. By R. McCabe, B. Ogera, H. Reeves. 2004. 
University Press of Colorado, 5589 Arapahoe Avenue, Suite 
206C, Boulder, Colorado 80303 USA. 208 pages. U.S. $29.95. 
Cloth. 


Snakes of the Americas: Checklist and Lexicon. By R. 
Tipton. Krieger Publishing, P.O. Box 9542, Melbourne, 
Florida 32902 USA. No price available. 


Status and Conservation of Florida Amphibians and Rep- 
tiles. By W. Meshaka, and K. Babbitt. 2005. Krieger Publishing 
P.O. Box 9542, Melbourne, Florida 32902 USA. No price 
available. 


*The Wolves of Algonquin Park a 12 Year Ecological 
study. By J. and M. Theberge. 2004. University of Waterloo, 
Waterloo, Ontario. Contact bkevans@fes.uwaterloo.ca. 
Canadian $ 23.50 Paper. 


Botany 


Biological Control of Invasive Plants in the United States. 
Edited by E. Coombs, J. Clark, G. Piper and A. Cofrancesco. 
2004. Oregon State University Press, 102 Adams Hall, Cor- 
vallis Oregon 97331 USA. 448 pages. U.S. $45.00. Paper. 


+ Emulating Natural Forest Landscape Disturbances — 
Concepts and Applications. By A. Perera, L. Buse, and M. 
Weber. 2004. Columbia University Press, 61 W. 62nd Street, 
New York, New York 10023 USA. 352 pages. U.S. $74.50. 
Cloth. 


*Gathering Moss — A Natural and Cultural History of 
Mosses. By R. Kimmerer. 2004. Oregon State University 
Press, 102 Adams Hall, Corvallis, Oregon 97331 USA. 176 
pages. U.S. $17.95. Paper. 


Klamath Heartlands — A guide to the Klamath Reserva- 
tion Forest Plan. By E. Wolf. 2004. Oregan State Universi- 
ty Press, 102 Adams Hall, Corvallis Oregon 97331 USA. 56 
pages. U.S. $19.95. Paper. 


2004 


Tree — A life Story. By D. Suzuki and W. Grady. 2004. Grey- 
stone, #201 — 2323 Quebec Street Vancouver, British Colum- 
bia V5T 4S7 Canada. 192 pages. Canadian $28.00 Cloth. 


Environment 


Death Takes a Gander. By Christine Goff. 2004. CLG, 
Box 3280, Evergreen, Colorado 80437 USA. 224 pages. 
U.S. $5.99. Paper. 


Managing Environmental Policy: A Casebook. By R. 
Watson, D. Kiel, and S. Robar. Krieger Publishing. P.O. 
Box 9542, Melbourne, Florida 32902 USA. 180 pages. U.S. 
$27.50. Cloth. 


Prairie — A Natural History. By Candace Savage. 2004. 
Greystone, #201 — 2323 Quebec Street Vancouver, British 


BOOK REVIEWS 


483 


Columbia VST 4S7 Canada. 320 pages. Canadian $60. 
Cloth. 


The Sacred Balance. By D. Suzuki and A. McConnnell. 
2004. Greystone #201 — 2323 Quebec Street, Vancouver, 
British Columbia V5T 4S7 Canada. 160 pages. Canadian 
$34.95. Cloth. 


Young Naturalists 


Hummingbirds. By L. Aziz. 2004. Firefly Books Ltd., 3680 
Victoria Park Avenue, Toronto, Ontario M2H 3K1 Canada. 
64 pages. Canadian $9.95. Paper. 


Penguins. By W. Lynch. 2004. Firefly Books Ltd., 3680 
Victoria Park Avenue, Toronto, Ontario M2H 3K1 Canada. 
64 pages. Canadian $ 9.95 Paper. 


484 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


News and Comment 


Froglog: Newsletter of the Declining Amphibian Populations Task Force (63, 64) 


Number 63, June 2004. Contents: Status of Three Species 
if Toads in North-western Mexico (Georgina Santos Barrera 
& Jesus Pacheco Rodriguez) — Ranids to the Rescue! (Jerry 
Lea & Luca Luiselli) — Effects of Habitat Loss and Frag- 
mentation on Anurans in Espinal Eco-region, Argentina: a 
GIS approach (Paola Peltzer, G. Bock, R. Tardivo & R. Laj- 
manovich) — DAPF Rapid Response Fund — Reports on 
DAPTF Seed Grants — Froglog Shorts. 

Number 64, August 2004. Contents: Morphological Abnor- 
malities in Frogs of West Java, Indonesia (Mirza D. Kusrini, 
Ross A. Alford, Anisa Fitri, Dede M. Nasir, Sumantri Rahard- 
yansah — Press Release: Concerns Remain About UV—B 
Damage To Amphibians (Andy Blaustein) — New Labora- 
tory of Natural Sounds in Cuba (Ansel Fong Grillo) — Reports 
on DAPTE Seed Grants (Tim Halliday) — WebFroglog [see 


Marine Turtle Newsletter (105) 


July 2004. 32 pages: articles: The Status and Conserva- 
tion of Sea Turtles in Kenya — Observations of Loggerhead 
Turtles Feeding on Discarded Fish Catch, Kefalonia — notes: 
Back to the Old Ways — letters to the editor: TEDs 00 — 
meeting reports — IUCNMTSG Update — book reviews — 
news & legal briefs — recent publications. 

The Marine Turtle Newsletter is edited by Brendan J. God- 
ley and Annette C. Broderick, Marine Turtle Research Group, 


www.opem.ac.uk/daptf and www.open.ac.uk/daptt/froglog] 
— Announcements and Meetings. 

Froglog is the bi-monthly newsletter of the Declining 
Amphibian Populations Task Force of The World Conservation 
Union (IUCN)/Species Survival Commission (SSC) and is 
supported by The Open University, The World Congress of 
Herpetology, and Arizona State University. The newsletter is 
Edited by John W. Wilkinson, Department of Biological Sci- 
ences, The Open University, Walton Hall, Milton Keynes, 
MK7 6AA, United Kingdom; e-mail: daptf@open.ac.uk. 
Funding for Froglog is underwritten by the Detroit Zoological 
Institute, P. O. Box 39, Royal Oak, Michigan 48068-0039, 
USA. Publication is also supported by Peace Frogs 
www.peacefrogs.com and by RANA and the US National 
Science Foundation grants DEB—0130273 and INT—0322375. 


Centre for Ecology and Conservation, University of Exeter 
in Cornwall, Tremough Campus, Penryn TR10 9EZ United 
Kingdom; e—mail MTN @seaturtle.org; Fax +44 1392 263700. 
Subscriptions and donations towards the production of the 
MTN can be made online at <http://www.seaturtle.org/mtn/> 
or postal mail to Michael Coyne (online Editor) Marine Tur- 
tle Newsletter, | Southampton Place, Durham, North Carolina 
27705 USA (e-mail: mcoyne @seaturtle.org). 


The Boreal Dip Net/L’Epuisette Boreal: Newsletter of the Canadian Amphibian and Reptile 
Conservation Network — Reseau Canadien de Conservation des Amphbiens et des Reptiles 


8(2) June 2004 


Contents: Editor’s Note — Kawartha Turtle Trauma Centre: 
Now a Recognized Charity — 2993 Digital Frog -CARCNET/ 
RECCAR Scholarship Winner Kimberley Pearson (Sara L. 
Ashpole) — A Survey if the Status of the Western Toad 
(Bufo boreas) in Mount Revelstoke and Glacier National 
Parks (Jesse Dykstra) — (Other) Upcoming Meetings: 18" 
Annual Meeting of the Society for Conservation Biology 
(Danna Schock) — The 9 Annual Meeting of the Canadian 
Amphibian and Reptile Conservation Network (CARCNET)/ 
Reseau Canadien de Conservation des Amphibiens et des 


Canadian Species at Risk May 2004 


Issued by the Committee on the Status of Endangered Wild- 
life in Canada (COSEWIC), the list is 49 pages containing: 
About COSEWIC: mandate, membership, definitions — 
Summary tables: Species designated in the “risk” and the 
Extinct categories and the Not at Risk, and Data Deficient 
categories (Tables 1-3) — Results of May 2004 COSEWIC 
meeting (Tables 4-5) — Explanation of status change sym- 
bols for reassessed species — COSEWIC Assessment Results 
— Table 6: Species assessed and designated Extinct: Extinct 
Category — Table 7: Species examined and designated in a 
“risk category”: Etripated Category, Endangered Category, 
Threatened Category, Special Concern Category — Table 8, 
Species assessed and found to be on the Not at Risk cate- 
gory — Table 9. Species considered and placed in the Data 


Reptiles RECCAR) Coast Terrace Inn, Edmonton, Alberta 
24-27, 2004 — Seasonal and Diurnal Patterns of Calling in 
Eastern Canadian Amphibians: Compiling the conventional 
wisdom as a null hypothesis (Frederick W. Schueler with con- 
tributions from Wayne F. Weller, John Gilhen, Lenny Shirose, 
and Brian Dalzell — Membership in CARCNET/RECCAR 
(contact Bruce Pauli, Canadian Wildlife Service, National 
Wildlife Research Centre, Carleton University, Raven Road, 
Ottawa, Ontario K1A OH3. Web site: http://www.carcnet.ca/). 


Deficient category — Record of Status Re-examinations — 
Record of Name Changes. Listed as of May 2004 (with 
November 2003 totals in parenthesis) are 12 (12) extinct, 21 
(21) extirpated, 169 (153) endangered, 114 (102) threatened, 
and 140 (143) of special concern. Of the 453 (431) forms in 
these categories, 66 (64) are mammals, 60 (56) birds, 32 
(31) reptiles, 19 (19) amphibians, 79 (807) fishes, 16 (13) 
arthropods, 21 (18) molluscs, 145 (136) vascular plants, 11 
(8) mosses, and 7 (6) lichens. In addition 151 (152) forms 
have been considered and found not at risk, and 33 (29) to 
be data deficient. 

This publication is available from COSEWIC Secretariat, 
Canadian Wildlife Service, Environment Canada, Ottawa, 
Ontario K1A 0H3. See Web site: http://www.cosewic.gc.ca. 


484 


2004 ERRATUM 485 


Erratum Canadian Field-Naturalist 118(2): 


Back cover contents omission: 


Fifteenth census of seabird populations in the sanctuaries of the North Shore of the Gulf 
of St. Lawrence 1998-1999 JEAN-FRANCOIS RAIL AND GILLES CHAPDELAINE 256 


Replacement Figure 2, page 161 for 
Morphology and population characteristics of Vancouver Island Cougars, Puma concolor vancouverensis, 
STEVEN F. WILSON, APRYL HAHN, AARON GLADDERS, KAREN M. L. Gon, AND DAviID M. SHACKLETON 


4 
@ Female 
3 & Male 
2 Males: 
y = 1.45Ln(x) - 0.54 
u R?= 0.70 
O | 
Ss 
6 0 
m Females: ® 
4 Md @ y = 0.52Ln(x) - 1.09 
R? = 0.34 
-2 
-3 
0 2 4 6 8 10 
Age (years) 


FIGURE 2. Relationship between body size (derived from the principal component analysis on morphological measurements, 
see text) and age for male and female Cougars in two Vancouver Island study areas. Logarithmic trend lines are 
presented for males (squares m) and females (diamonds ®) pooled by study area. The body size measurement was 
used because it was less condition-dependent than using total weight alone. 


Minutes of the 125 Annual Business Meeting of 
The Ottawa Field-Naturalists’ Club 13 January 2004 


Place and time: Canadian Museum of Nature, Ottawa, Ontario, 7:30 p.m. 


Garry McNulty, President 
Thirty-five persons attended the meeting. 


Chairperson: 
Attendance: 


Attendees spent the first half-hour reviewing the minutes of the previous meeting, the Treasurer’s Report and the Report of 
Council. The meeting was called to order at 7:35 p.m. with some opening remarks from Gary McNulty, the President. 


1. Minutes of the Previous Meeting 
There were no changes to the minutes of the 124" Annual 
Business Meeting. 
It was moved by Fenja Brodo/David Hobden that the min- 
utes be accepted. 
(Motion Carried) 


2. Business Arising from the Minutes 

Garry McNulty inquired whether we were still receiving 
the annual reports for the Trinidad and Tobago Naturalists’ 
Club. Frank Pope reported that we were receiving their annual 
reports along with their newsletters. 

The Soiree was revamped last year. Changes were made 
to the venue and entertainment was included in the course of 
the evening. Fenja Brodo, the new chair of the Excursions and 
Lectures Committee, will be working with the Education and 
Publicity Committee to continue the process of making the 
Soiree an evening that members will not want to miss. 


3. Communications Relating to the Annual 
Business Meeting 
There were no communications relating to the Annual 
Business Meeting. 


4. Treasurer’s Report 
Frank Pope reviewed the financial report for the year end- 
ing September 30, 2003, noting that the Club’s net assets had 
decreased by approximately $60,000. This was due in large 
part to the contribution of $55,000 to the Alfred Bog Fund. 
Moved by Frank Pope and seconded by Roy John that the 


Financial Report be accepted. 
(Motion Carried) 


5. Committee Reports 

Gary McNulty introduced each of the Committee reports 
and a representative of the appropriate Committee and asked 
for questions and comments. He thanked the committee chairs 
and committee members for their work over the past year. 

Moved by Mike Murphy/Eleanor Zurbrigg, that the reports 
be accepted. 

(Motion Carried) 


6. Nomination of the Auditor 
Moved by Frank Pope, seconded by Bill Cody, that Janet 
Gehr continue as Auditor for another year. 
(Motion Carried) 


7. Report of the Nominating Committee 


President Mike Murphy 
Vice President Gillian Marston 
Secretary Susan Laurie Bourque 
Treasurer Frank Pope 
Past President Gary McNulty 
Business Manager Bill Cody 
Editor CFN Francis Cook 
Editor T&L Karen McLachlan Hamilton 
Committee Chairs 

Birds Chris Traynor 
Computers 

Conservation Stan Rosenbaum 
E&P John Cameron 
E&L Fenja Brodo 
Finance Louise Schwartz 
FWG David Hobden 
Macoun rep Barbara Gaertner 
Membership Dave Smythe 
Publications Ron Bedford 
FON Rep Cendrine Huemer 
Members at large 

Kathy Conlan 

Diane Lepage 


Barbara Gaertner 
Christine Lewis 
Henry Steger 


Chairs not on Council 


Awards Ernie Brodo 

Macoun Rob Lee 

Nominations Fenja Brodo 

Retiring from the Council: = Charles Clifford, Roy John, 
Marcel Gahbauer, 


Eleanor Zurbrigg 


New on the Council: Christina Lewis, Henry Steger 
Moved by Fenja Brodo, seconded by Frank Pope, that the 

slate of nominations for the 2004 Council be accepted. 
(Motion Carried) 


8. New Business 
There was no new business. 


9. Presentation by David Hobden, ““FWG 
and its Rise to Fame” 
David Hobden presented an enjoyable and informative 
digital photographic presentation created by Henry Steger and 


486 


2004 


David Hobden. The presentation touched on the develop- 
ment and flora and fauna of the Fletcher Wildlife Garden. 
The digital format successfully combined prints, slides and 
digital pictures. It was very well received. 


MINUTES OF THE 125™ ANNUAL BUSINESS MEETING 


487 


10. Adjournment 

Fenja Brodo made an announcement about the upcoming 
4"" Annual Great Backdoor Bird Count. 

Moved by Fenja Brodo/David Hobden that the meeting 
be adjourned at 9:15 pm. (Motion Carried) 


SUSAN LAURIE-BOURQUE, Recording Secretary 


The Ottawa Field-Naturalists’ Club Committee Reports for 2003 


Awards Committee 

The Awards Committee met in January to consider nomi- 
nations made for the various OFNC Awards. As a result of 
our deliberations, the following four awards for the year 2002 
were presented at the OFNC’s Annual Soirée, which took 
place 26 April 2003 at St. Basil’s Church in Ottawa. 


MEMBER OF THE YEAR: 

Bill Royds: For his many contributions to the Club’s con- 
servation efforts, especially during this last year as the link 
between the OFNC and the Greenspace Alliance of Canada’s 
Capital (GACC). 


GEORGE MCGEE SERVICE AWARD: 

David Hobden: For his active participation in the Con- 
servation Committee and his enthusiastic, productive work 
on the Fletcher Wildlife Garden Committee. 


CONSERVATION AWARD — MEMBER: 

Philip Fry: For establishing and maintaining the “Old 
Field Garden” near Oxford Mills as a model for habitat 
restoration and wildflower gardening. 


CONSERVATION AWARD — NON-MEMBER: 

Michéle André-St. Cyr: For her relentless work in pre- 
venting the death of turtles along the highways of southern 
Ontario by the establishment of “Turtle Crossing” signs, 
and for her active and successful efforts in educating the 
public about turtles. 

I. BRoDO 


Birds Committee 
The Birds Committee participated jointly with the Club des 
Ornithologues de l’Outaouais to run a successful Christmas 
Bird Count in December 2002. The fall bird count was held 
again in October 2003 and continues to grow in popularity. 
We were again active in the Peregrine Falcon Watch at the 
downtown nest site. One female was successfully fledged. 
The Bird Record Sub-committee continues to review records 
of rare birds and is currently compiling a lengthy database 
of Ottawa bird records. The Bird Record Sub-committee 
also finished publication of the new Ottawa District Bird 
Checklist 2002. The checklist is available in many Ottawa 
nature stores. The third year of surveying for the Ontario 
Breeding Bird Atlas (Region 24) has been completed. There 
are plans to add five more under-surveyed squares from an 
adjacent region for the 2004 season. This years seed-a-thon 
raised close to $400.00 for the club’s bird feeders. We con- 
tinue to operate the rare bird alert and the Ottawa bird status 
line, a recorded telephone message of current bird sightings. 
Some members of the Committee participated in the Taver- 

ner Cup competition 
C. TRAYNOR 


Computer Management Committee 
This year, the Committee reviewed and revised their Terms 
of Reference and the revisions were approved by the Council. 


It also reviewed the state of the Club’s Membership Data- 
base System application. Upon a recommendation by the 
Computer Management Committee, the Council approved 
the acquisition of Microsoft Access as the Club’s data entry 
and database management technology. This product provides 
a relational database engine with VisualBasic language sup- 
port for applications development. The Membership Database 
application will be redeveloped using Microsoft Access 
during 2004. 

M. MurpHy 


Conservation Committee 

Since November 2002 the CC has been assisting local 
residents calling themselves Larose Defense Fund/Fond de 
défense Larose with opposition to a proposed zoning change 
within part of the forest. We expressed support for the preser- 
vation of francophone heritage at other locations. The Club 
contributed $1000 towards the Fund’s legal costs at an OMB 
hearing. 

In cooperation with the Greenspace Alliance of Canada’s 
Capital, the CC appeared at City Planning and Development 
Committee in regard to the 3rd draft of the Official Plan. 
Some improvements were secured. 

We helped the FON to hold this workshop at the Museum 
of Nature in Ottawa. 

Several Club members reviewed the City’s list of areas to 
be studied and suggested additional areas. Adrianne Sinclair 
of the CC was appointed to the Public Advisory Committee. 

What in the 1970s the OFNC called the South Gloucester 
Conservation Area gradually became a quarry, but the 177 
City-owned acres were known to support uncommon plant 
species. Despite intervention by the CC and the Greenspace 
Alliance, the City voted to sell this land to the quarry owner 
for $1.7M, but imposed requirements for environmental 
assessment, and assigned the sale revenue to a natural land 
acquisition fund. 

S. ROSENBAUM 


Education And Publicity Committee 

The committee organized and managed OFNC participa- 
tion in three major public events: the Ottawa Teachers Fed- 
eration professional development day, the Carlingwood Wild- 
life Festival and Health Canada’s celebration of Environment 
Week. The Wildlife Festival was the largest and most success- 
ful in attracting public interest. Twenty OFNC volunteers and 
five Macoun members assisted with this event. 

A demonstration of bird songs using tapes and slides was 
provided as a pre-concert activity for the Young Peoples Con- 
cert Series at the NAC, April 26. 

The Ottawa Field-Naturalists’ Club presented three Natural 
History Awards of $50.00 each plus an OFNC membership 
at the Ottawa Regional Science Fair. The judges were Kathy 
Conlan and Susan Laurie-Bourque. 

A French version of the club brochure was prepared and 
production begun at year’s end. 


488 


The sales table at the club’s evening presentations at the 
Canadian Museum of Nature provided revenue of $400. 
J. CAMERON 


Excursions And Lectures Committee 

In 2003 this Committee arranged 38 events and nine 
monthly meetings including the ABM and the Soirée. Seven 
general interest excursions were offered to Club members. 
The Birds Committee ran 12 birding excursions, three bird 
counts as well as assisted with The Taverner Cup. Two of our 
monthly meetings were also devoted to birds. There were 
six botanical excursions or workshops and two mushroom 
activities. Insects were the subject of one excursion and two 
of our monthly meetings. Other excursions were for mudpup- 
pies, astronomy and a museum visit. Workshops were also 
given on geology and on digital cameras. Two of our monthly 
meetings were devoted to conservation matters and one intro- 
duced us to polar environments. 

The monthly meetings drew about 50 people on average. 
Excursions and workshops are designed for a smaller number 
of participants. This year we asked leaders to have partici- 
pants sign a waiver form. The form is primarily designed to 
inform participants about liability but it provides a record of 
attendance. 

F. BRODO 


Executive Committee 

The Executive Committee met in October to review sey- 
eral items that required responses and/or action plans. As a 
result of that meeting, action plans have been put in place to 
address the following issues in 2004: 

(1) review of electronic publication possibilities 

(2) digital photo archives 

(3) how to retain new members 

(4) system to record names and number of persons parti- 
cipating in club excursions and activities. 

The Committee also reviewed the club’s liability insurance, 
its role in the Taverner Cup, the issue of attracting volunteers 
and it put in place a succession planning exercise and a sys- 
tem to review and report on the publications of other nature 
clubs. These matters require further action in 2004 but 
should be completed by year end. 

G. McNuLTy 


Finance Committee 
The Finance Committee met three times in calendar year 

2003. The following items are highlighted: — a review of 

existing and proposed honoraria was undertaken. No changes 

to the current structure were recommended to Council. 

— the committee recommended to Council that the Alfred 
Bog Fund be topped up by $4,000 from the de Kiriline 
Lawrence Fund, bringing the total OFNC contribution to 
$59,000. 

— options for replacement accounting software were assessed 
but no decision was taken. 

— areview of membership fees for 2004 was carried out, and 
the Committee recommended to Council that the fees not 
be raised for the upcoming year. However, given the ex- 
pected year-end deficit, this issue will need serious con- 
sideration next year. 

— a budget for fiscal year 2003-04 was tabled and approved 
at Council in the fall, showing a small projected deficit. 

— other items such as the RoweCom bankruptcy, liability 
and directors’ insurance, and Taverner Cup funding were 
discussed. 

L. SCHWARTZ 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Fletcher Wildlife Garden 

Volunteers contributed over 3000 hours, mainly on Friday 
or Sunday mornings. Their work is particularly noticeable 
in the Backyard Garden. The Management Committee met 
eleven times during the year. 

The big achievement for the year was the production of 
an agreement with Agricultureand Agri-Food Canada for 
use of the site and occupation of the building. Negotiations 
were completed in the summer and the agreement finally 
signed in November. It is to run until April 2006. We wish 
to thank Frank Pope for handling legal and insurance issues 
and the final signing. 

During the year we held a public event for International 
Migratory Bird Day in May and our best plant sale ever in 
June thanks to Algonquin College and our own volunteers. 

Our traveling display was seen at Seedy Saturday, The 
Wildlife Festival, Environment Day at Health Canada, an 
Invasive Species Workshop at the Museum of Nature and at 
the OFNC Soiree. 

This year we had a part-time summer employee, since 
we did not obtain a grant to support the position as we had 
in previous years. She dealt with visitors, kept the Interpre- 
tation Centre open and worked mainly on invasive plant con- 
trol, supplementing the work of a group of volunteers. 

The Interpretation Centre was opened by volunteers on 
Sunday afternoons through the summer. Located there is a 
small library, including a complete collection of the Cana- 
dian Field-Naturalist, and two stereo microscopes for examin- 
ing small specimens. The Centre was used for meetings by 
OFNC Council and four committees and for workshops. 

D. HOBDEN 


Macoun Field Club Committee 

The committee met just once, to plan the start-up of the 
Macoun Club year; much of the work of organizing the pro- 
gram from month to month was done over the telephone and 
by e-mail. Some committee members arranged for speakers 
each week and supervised the meetings, while others led 
field trips twice a month. Some also led the high-school-age 
members on camping trips at Christmas and in June. 

Through this program, the children and teenagers of the 
Macoun Club were introduced to local geology, lichens, plant 
biology, insects, and wildlife (both Canadian and exotic). Field 
trips gave members hands-on experience in identifying trees 
and animal tracks, and allowed them to take part in the Club’s 
ongoing research on butternut canker disease, lichen bio- 
diversity, and porcupine populations. 

The monthly newsletter and the annually produced Little 
Bear magazine were both prepared by the young people them- 
selves, while the Club’s web site was maintained jointly with 
Committee members. Individual Macoun Club members pre- 
sented natural science projects at the OFNC Soirée in April, 
and the Club’s executive recounted the group’s activities for 
the OFNC members assembled there. 

R. LEE 


Membership Committee 
The distribution of memberships for 2003 is shown in the 
table (below), with the comparable numbers for 2002 in 
brackets. These statistics do not include the 23 affiliate orga- 
nizations which receive free copies of the Club’s publications. 
This year, the Club lost a long time member and valuable 
contributor to the production of the Canadian Field-Naturalist 
(CEN), with the death of Dr. Wilson Eedy, member since 
1970 and Book-Review Editor of the CFN for many years. 
D. SMYTHE 


2004 MINUTES OF THE 125™ ANNUAL BUSINESS MEETING 489 

| ° . . sae . 

| Distribution of Memberships in The Ottawa Field-Naturalists’ Club 

CANADIAN FOREIGN 

| Type Local Other USA Other Total 

_ Family 317 (340) 28 (29) | (1) | (2) 342 (372) 
Individual 323 (345) Le C20) 24 (26) 6 (5) 464 (496) 
Honorary 15 (15) 9 (9) 0 (0) 0 (0) 24 (24) 
Life 21 (21) 20 (21) 7 (5S) | (1) 49 (48) 
Sustaining 11 (8) 3 (2) 0 (O) 0) (QO) 1] (10) 
Total 687 (729) 166 (181) 32. (32) 8 (8) 890 (950) 


Publications Committee 

The Publications Committee met twice in 2003. 

_ Four issues of The Canadian Field-Naturalist were pub- 
— lished in 2003: Volume 116, #2,3,4 and Volume 117, #1. 
These four issues contained 698 pages; 47 articles; 20 notes; 
_ 1 COSEWIC article; 81 book reviews; 144 new titles; 2 
_ commemorative tributes; 24 pages of News and Comments; 
and a 29 page index. 

The long-time Book Review Editor, Wilson Eedy, died sud- 
denly in the summer. His position has been taken over by 
Roy John. All of the Associate Editors will be reappointed 
for 2004. 

Six articles qualified for support from the Manning Mem- 
_ orial Fund for a total of $4044. 


Steady progress was made in getting the publication sched- 
ule back on track. The printing of The CFN has been taken 
from St. Joseph and contracted to Gilmore Printing Services 
Inc. beginning with Volume 117, #2. It is expected that this 
will both reduce the printing costs and improve the photo- 
graphic quality. 

In response to an author’s complaint, the details of pub- 
lication charges have been reworded. The new text says, in 
effect, that only members of The Ottawa Field-Naturalists’ 
Club and subscribers to The CFN are eligible to apply for 
waiver of page charges for the first five pages, and then only 
if institutional funds are unavailable. 

Volume 37 of Trail & Landscape was published in four 
issues containing 192 pages with the usual mix of good articles 
and news of Club events. 

R. BEDFORD 


490 


Auditor’s Report 


THE CANADIAN FIELD-NATURALIST 


To The Members of THE OTTAWA FIELD NATURALISTS’ CLUB 


I have audited the balance sheet of THE OTTAWA 
FIELD-NATURALISTS’ Club as at September 30, 
2003, the statement of changes in net assets, and the 
statements of operations. These financial statements are 
the responsibility of the organization’s management. 
My responsibility is to express an opinion on these 
statements based on my audit. 

Except as explained in the following paragraph, I 
conducted my audit in accordance with generally ac- 
cepted auditing standards. Those standards require that 
I plan and perform an audit to obtain reasonable as- 
surance whether the financial statements are free of 
material misstatement. An audit includes examining 
evidence supporting the amounts and disclosures in 
the financial statements. An audit also includes assess- 
ing the accounting principles used and significant esti- 
mates made by management, as well as evaluating the 
overall financial statement presentation. 

In common with many non-profit organizations, 
the Ottawa Field-Naturalists’ Club derives some of its 
revenue from memberships, donations, and fund raising 
activities. These revenues are not readily susceptible to 
complete audit verification, and accordingly, my veri- 
fication was limited to accounting for the amounts 
reflected in the records of the organization. 

In my opinion, except for the effect of the adjust- 
ments, if any, which I might have determined to be 
necessary had I been able to satisfy myself concern- 
ing the completeness of the revenues referred to in 
the preceding paragraph, these financial statements 
present fairly, in all material respects, the financial 
position of the OFNC as at September 30, 2003, and 
the results of its operations and changes in net assets 
for the year then ended in accordance with generally 
accepted accounting principles. 


JANET M. GEHR 
Chartered Accountant 
North Gower, Ontario 
January 12, 2004 


Vol. 118 
The Ottawa Field-Naturalists’ Club 
Balance Sheet 
September 30, 2003 
2003 2002 
ASSETS 
CURRENT 
Cash (Note 1) $ 2,781 $ 71,087 
Investment certificates (Note 1) 43,764 84,790 
Marketable securities (Note 2) 246,467 235,758 
Accounts receivable 27,690 7,238 
Prepaid expenses 1,000 1,000 
321,702 399,873 
CAPITAL ASSETS (Note 3) - _ 
Land — Alfred Bog 3,348 3,348 
$ 325,050 $403,221 
LIABILITIES AND FUND BALANCES 
CURRENT 
Accounts payable and 
accrued liabilities $ 2,000 $ 2,000 
Deferred revenue 12,075 13,500 
14,075 15,500 
Life memberships 12,663 2A 
NET ASSETS 
Unrestricted 60,609 80,325 
Club reserve 100,000 100,000 
Manning principal 100,000 100,000 
Manning interest - OFNC 1,434 1,963 
- CFN 13,153 13,179 
Seedathon 1,618 2,087 
Anne Hanes memorial 870 870 
de Kiriline-Lawrence 18,798 23,975 
Macoun Baillie Birdathon 3 5857/ S27 
Alfred Bog 493 52,274 
298,312 376,000 
$ 325,050 $403,221 


2004 MINUTES OF THE 125" ANNUAL BUSINESS MEETING 49] 


The Ottawa Field-Naturalists’ Club The Ottawa Field-Naturalists’ Club 
Statement of Operations Canadian Field-Naturalists — Statement Of Operations 
For the Year Ended September 30, 2003 For the Year Ended September 30, 2003 
2003 2002 2003 2002 
REVENUE REVENUE 
Memberships $ 14,615 $ 15,488 Memberships $ 9,743 $ 10,325 
Trail and Landscape 193 196 Subscriptions 25,369 28,483 
Interest 1,339 1,457 Reprints 9,534 5,608 
GST rebate 780 4,532 Publication charges 50,559 16,812 
Other 617 760 Interest and exchange 7,940 10,404 
17,547 22,433 GST rebate 3,040 3,448 
SS a Other 734 339 
OPERATING EXPENSES 106,919 75,419 
Affiliation fees 680 680 
Computer 302 216 EXPENSES 
Membership Syl 1,437 Publishing $ 80,758 $ 61,570 
Office Assistant 1,000 1,000 Reprints 7,307 4,127 
Telephone 1,622 1,848 Circulation 11,307 7,653 
Insurance 655 655 Editing 4,183 2,652 
Audit 1,000 1,000 Office Assistant 5,000 5,000 
GST 1,290 136 Honoraria 9,000 9,000 
Other 884 2,847 GST rebate 6,508 5,146 
Tal RE: Suet 124,615 95,569 
CLusB ACTIVITY EXPENSES ExcEss EXPENSES OVER 
Awards is 471 REVENUE $ (17,696) $ (20,150) 
Birds 1,696 223 ae Saupe 
Education and Publicity 73 1322 
Excursions and lectures 496 164 
Macoun Field Club 851 1,027 
Soiree 19 466 
Trail and Landscape 8,368 9,088 
Fletcher Wildlife Garden (Note 4) 2,623 1,294 
Other = _ 
14,126 14,055 
22,710 25,054 


Excess EXPENSES OVER 
REVENUE $ (5,163) $ (2,621) 


492 THE CANADIAN FIELD-NATURALIST 


The Ottawa Field-Naturalists’ Club Notes to the Financial Statements 
September 30, 2003 


1. CASH 


Chequing 

Savings 

Nesbitt Burns 

Fletcher Wildlife Garden 


Investment Certificates: Maturity Maturity 
Value Date 
$ 44,385 04/29/04 


2. MARKETABLE SECURITIES 


Investment Certificates: Maturity Maturity 
Value Date 
Province of Newfoundland Coupon $ 44,782 10/17/11 
Province of Manitoba Coupon 29,847 11/15/04 
Ontario Savings Bonds 40,000 06/21/05 
CMHC Global Debs 52,000 12/01/06 
Province of Ontario Bond 30,000 09/12/07 
Province of Newfoundland Bond 20,000 10/07/08 
Government of Canada Coupon 30,167 12/01/09 
Province of New Brunswick Bond 20,000 06/15/10 


3. CAPITAL ASSETS 
Equipment at a cost of $16,748 is fully amortized. 


4. FLETCHER WILDLIFE GARDEN 


REVENUE 
Federal government 
City of Ottawa 
Taverner Cup 
Sales 
GST 
Donations 
Other 


EXPENSES 
Program 

Backyard 

Habitats 
Interpretation centre 
Administration 
Publications 

GST 

Library 

Other 


2003 


$ (3,777) 
1,344 

919 
4,295 


$ 2,781 
Yield 


1.00% 


Yield 


4.525% 
5.119% 
6.45% 

5.250% 
6.125% 
6.263% 
5.605% 
6.231% 


243 
132 


6,346 
$ (2,623) 


Vol. 118 ! 


2002 


$ 5,589 
58,336 
919 
5,868 


$ 70,712 


Book 
Value 
$ 43,764 


Book 
Value 

$ 31,421 
26,754 
41,370 
53,539 
31,187 
20,538 
20,590 
21,068 


$246,467 


2002 


$ 3,059 
1,450 
1,354 

872 
398 
292 


7,425 


3,174 
123 
1,864 
783 
1,048 
337 
303 
87 


8,719 
$ (1,294) 


2004 MINUTES OF THE 125™ ANNUAL BUSINESS MEETING 493 
The Ottawa Field-Naturalists’ Club Statement of Changes in Net Assets 
For the Year Ended September 30, 2003 (Note 5) 
Net Beginning Excess Excess Other Ending 
Assets Balance Expenses CFN Expenses OFNC Revenue Expenses Balance 
Unrestricted $ 80,325 $ (17,696) $ (5,163) $ 3,143 a $ 60,609 
Club Reserve 100,000 - - - ~ 100,000 
Manning Principal 100,000 = - - - 100,000 
Manning — OFNC (a) 1,963 ~ - 1,021 1,550 1,434 
Manning — CEN (b) 13,179 ~ ~ 4,086 4,112 13,153 
Seedathon 2,087 - - 502 971 1,618 
Anne Hanes Memorial 870 - - = ~ 870 
de Kiriline-Lawrence (c) 23,975 _ - 381 5,558 18,798 
Macoun Baillie Birdathon (d) 1.327 — - 10 - 1,337 
Alfred Bog (e) 52,274 - - 3,219 55,000 493 
$ 376,000 $ (17,696) $ (5,163) $ 12,362 $ 67,191 $ 298,312 


STATEMENT OF CHANGES IN NET ASSETS 


a) Manning OFNC Expenses: Taverner Competition, $1,250; Peregrine Watch, $300; 

b) Manning CFN Expenses: Vol. 114, No. 2, Douglas Clark, Grizzly Bears in Northern Manitoba $228; Vol. 116, No. 1, 
Mark Hipfner et al., Common Eiders around Digges Sound $340; Gerald Kuzyk et al., Wolf response to sled dog 
$160; Kevin White et al., Predation of wolves on wolverines and martens $160; W. A. Fuller, Canada and the buffalo 
$1,609; Vol. 116, No. 3, John Goold et al., Sperm whale strandings $1,615. 

c) de Kiriline Lawrence Expenses: Nature Conservancy of Canada — land in Alfred Bog $4,000; Sierra Club, Ottawa 
Group — Leitrim Wetlands publicity — $1,000; Greenspace Alliance (Unger) — newspaper ad in defence of South 


Gloucester Conservation Area $558. 


d) Alfred Bog Expense: Nature Conservancy of Canada, $55,000 proceeds from fund raising campaign for acquisition 


of land in Alfred Bog. 


The Ottawa Field-Naturalists’ Club Summary of Significant Accounting Policies 


September 30, 2003 
1. Nature of Business 

The organization is non-profit and incorporated under the 
laws of Ontario (1884). The organization promotes the appre- 
ciation, preservation, and conservation of Canada’s natural 
heritage. It encourages investigation and publishes the results 
of the research in all fields of natural history and diffuses 
information on these fields as widely as possible. It also sup- 
ports and cooperates with other organizations engaging in 
preserving, maintaining or restoring environments of high 
quality for living things. 
2. Financial Instruments 

The organization’s financial instruments consist of cash, 
accounts receivable, marketable securities, and accounts pay- 
able. Unless otherwise noted, it is the management’s opinion 
that the organization is not exposed to significant interest, 
currency, or credit risks arising from these financial instru- 
ments. The fair value of these instruments approximate their 
carrying values. 


3. Capital Assets 

Capital assets acquired after 1989 are expenses. Capital 
assets acquired prior to 1990 were recorded as assets at cost 
and amortized on a straight-line basis. These assets have 
been fully amortized. 
4. Statement of Changes in Financial Position 

A statement of changes in financial position has not been 
provided as it would not provide additional meaningful infor- 
mation. 
5. Foreign Currency 

Transactions during the year in U.S. dollars have been 
converted in the accounts to Canadian dollars at the exchange 
rate effective at the date of the transaction. All monetary assets 
in U.S. dollars at year end have been converted to Canadian 
dollars at the rate effective on Sept. 30, 2003. Gains or losses 
resulting therefrom are included in revenue or expenses. 


Advice for Contributors to The Canadian Field-Naturalist 


Content 

The Canadian Field-Naturalist is a medium for the publi- 
cation of scientific papers by amateur and professional natu- 
ralists or field-biologists reporting observations and results 
of investigations in any field of natural history provided that 
they are original, significant, and relevant to Canada. All read- 
ers and other potential contributors are invited to submit for 
consideration their manuscripts meeting these criteria. The 
journal also publishes natural history news and comment items 
if judged by the Editor to be of interest to readers and sub- 
scribers, and book reviews. Please correspond with the Book 
Review Editor concerning suitability of manuscripts for this 
section. For further information consult: A Publication Policy 
for the Ottawa Field-Naturalists’ Club, 1983. The Canadian 
Field-Naturalist 97(2): 231-234. Potential contributors who 
are neither members of The Ottawa Field-Naturalists’ Club 
nor subscribers to The Canadian Field-Naturalist are encour- 
aged to support the journal by becoming either members or 
subscribers. 


Manuscripts 

Please submit, to the Editor, in either English or French, 
three complete manuscripts written in the journal style. 
The research reported should be original. It is recommended 
that authors ask qualified persons to appraise the paper before 
it is submitted. AJ] authors should have read and approved it. 
Institutional or contract approval for the publication of the data 
must have been obtained by the authors. Also authors are ex- 
pected to have complied with all pertinent legislation regard- 
ing the study, disturbance, or collection of animals, plants or 
minerals. The place where voucher specimens have been de- 
posited, and their catalogue numbers, should be given. Lati- 
tude and longitude should be included for all individual local- 
ities where collections or observations have been made. 

Print the manuscript on standard-size paper, doublespace 
throughout, leave generous margins to allow for copy mark- 
ing, and number each page. For Articles and Notes provide 
a citation strip, an abstract and a list of key words. Gener- 
ally, words should not be abbreviated but use SI symbols for 
units of measure. The names of authors of scientific names may 
be omitted except in taxonomic manuscripts or other papers 
involving nomenclatural problems. “Standard” common names 
(with initial letters capitalized) should be used at least once 
for all species of higher animals and plants; all should also 
be identified by scientific name. 

The names of journals in the Literature Cited should be 
written out in full. Unpublished reports and web documents 
should not be cited here but placed in the text or in a sepa- 
rate Documents Cited section. List the captions for figures 
numbered in arabic numerals and typed together on a separate 
page. Present the tables each titled, numbered consecutively 
in arabic numerals, and placed on a separate page. Mark in 
the margin of the text the places for the figures and tables. 


Check recent issues (particularly Literature Cited) for 
journal format. Either “British” or “American” spellings are 
acceptable in English but should be consistent within one 
manuscript. The Oxford English Dictionary, Webster’s 
New International Dictionary and le Grand Larousse 
Encyclopédique are the authorities for spelling. 


Illustrations 

Photographs should have a glossy finish and show sharp 
contrasts. Electronic versions should be high resolution. Photo- 
graphic reproduction of line drawings, no larger than a 
standard page, are preferable to large originals. Prepare line 
drawings with India ink on good quality paper and letter (don’t 
type) descriptive matter. Write author’s name, title of paper, and 
figure number on the lower left corner or on the back of each 
illustration. 


Reviewing Policy 

Manuscripts submitted to The Canadian Field-Naturalist 
are normally sent for evaluation to an Associate Editor (who 
reviews it or asks another qualified person to do so), and at 
least one other reviewer, who is a specialist in the field, cho- 
sen by the Editor. Authors are encouraged to suggest names 
of suitable referees. Reviewers are asked to give a general 
appraisal of the manuscript followed by specific comments 
and constructive recommendations. Almost all manuscripts 
accepted for publication have undergone revision—sometimes 
extensive revision and reappraisal. The Editor makes the 
final decision on whether a manuscript is acceptable for pub- 
lication, and in so doing aims to maintain the scientific quality, 
content, overall high standards and consistency of style, of 
the joumal. 


Special Charges — Please take note 

Authors must share in the cost of publication by paying 
$80 for each page, plus $15 for each illustration (any size up 
to a full page), and up to $80 per page for tables (depending 
on size). Authors may also be charged for their changes in 
proofs. Reproduction of color photos is extremely expensive; 
price quotations may be obtained from the Business Manager. 
If grant or institutional funds are not available, club members 
and subscribers may apply for a waiver of charges for the 
first five pages. 

Limited joumal funds are available to help offset publi- 
cation charges to authors with minimal financial resources. 
Requests for financial assistance should be made to the Busi- 
ness Manager when the manuscript is accepted. 


Reprints 
An order form for the purchase of repents will accompany 
the galley proofs sent to the authors. 


FRANCIS R. Cook, Editor 
RR 3 North Augusta, Ontario KOG IRO Canada 


494 


——SaEaS==—_—_=>—=—>= 


TABLE OF CONTENTS (concluded) Volume 118 Number 3 


} 
ik 


Sequential polyandry in Piping Plover, Charadrius melodus, nesting in eastern Canada 
DIANE L. AMIRAULT, JONATHAN KIERSTEAD, PETER MACDONALD, and LARRY MACDONNELL 


| American Dipper, Cinclus mexicanus, preys upon larval Tailed Frogs, Ascaphus truei 
CurisTy A. Morrissey and ROBERTA OLENICK 


Piping Plover, Charadrius melodus, egg viability after seawater immersion 
JULIE MCKNIGHT, LINDA THOMAS, and DIANE L. AMIRAULT 


Tribute 
1A tribute to Loris Shano Russell, 1904-1998 KEVIN L. SEYMOUR 


|Book Reviews 

|ZooLocy: The Uses and Curation of Bird’s Egg Collections: An Examination and Bibliography — Birds of 
| Nunavut: A Checklist — Birds of the Untamed West: The History of Birdlife in Nebraska, 1750 to 1875 — 
Birds of Nebraska: Their Distribution and Temporal Occurrence — Canadian Skin and Scales: A Complete 
Encyclopedia of Canadian Amphibians and Reptiles — Conservation and Ecology of Turtles of the Mid- 
Atlantic Region: A Symposium — For Love of Insects — The Freshwater Fishes of Manitoba — The 
History of Ornithology in Virginia — All-Weather Hawk Watcher’s Field Journal —- Self-Portrait with 
Turtles: A Memoir — Wild Mammals of North America: Biology, Management, and Conservation 
(Second edition) — Pete Dunne on Bird Watching: The How-to, Where-to and When-to of Birding 


-|Borany: Alpine Plants of North America: An Encyclopaedia of Mountain Flowers from Rockies to Alaska 
— Arboretum America, a Philosophy of the Forest — Cape Cod Wiidflowers: A Vanishing Heritage — 
Lewis Clark’s Field Guide to Wild Flowers of the Sea Coast in the Pacific Northwest — Wild Flowers of 
Field & Slope in the Pacific Northwest — Wild Flowers of Forest & Woodland in the Pacific Northwest 
Wild Flowers of the Mountains of the Pacific Northwest 


ENVIRONMENT: Conserving Living Natural Resources in the Context of a Changing World — Visions of the 
Land: Science Literature, and the American Environment from the Era of Exploration to the Age of 
Ecology — Genetic, Demography and Viability of Fragmented Populations — A Primer of Ecological 
Genetics 


MISCELLANEOUS: Smithsonian Institution Secretary Charles Doolittle Walcott 


NEw TITLES 


|News and Comment 

Froglog: Newsletter of the Declining Amphibian Populations Task Force (63, 64) — Marine Turtle 
Newsletter (105) — The Boreal Dip Net/L’Epuisette Boreal: Newsletter of the Canadian Amphibian and 
Reptile Conservation Network - Reseau Canadien de Conservation des Amphbiens et des Reptiles 8(2) 
June 2004 — Canadian Species at Risk May 2004 


_|Errata: Canadian Field-Naturalist 118(2) 
Omission from the back cover contents: Rail and Chapdelaine 
Replacement figure for Wilson et al. page 161 


Minutes of the 125 Annual Business Meeting of The Ottawa Field-Naturalists’ Club 
Tuesday 13 January 2004 


| Advise to Contributors to The Canadian Field-Naturalist 


Mailing date of the previous issue 118(2): 10 May 2005 


2004 


444 


446 


448 


45] 


465 


474 


477 
481 
483 


484 


485 


486 


494 


THE CANADIAN FIELD-NATURALIST Volume 118 Number 3 


Articles 


Population dynamics of Deer Mice, Peromyscus maniculatus, and Yellow-pine 
Chipmunks, Zamias amoenus, in old field and orchard habitats 
THOMAS P. SULLIVAN, DRUSCILLA S. SULLIVAN, and EUGENE J. HOGUE 


Density and survival of lady beetles (Coccinellidae) in overwintering sites in Manitoba 
W. J. TURNOCK and I. L. WISE 


Description of age-O Round Goby, Neogobius melanostomus Pallus (Gobiidae) and ecotone 
utilization in St. Clair lowland waters, Ontario JOHN K. LESLIE and CHARLES A. TIMMINS 


A systematic analysis of the alpine saxifrage complex (Saxifragaceae) in the Canadian Arctic Islands 
using morphology and chloroplast DNA data CAROLINE HEALY and LYNN J. GILLESPIE 


The influence of prey availability and vegetation characteristics on scent station visitation rates 
of Coyotes in a heterogeneous environment LYNDA A. RANDA and JOHN A. YUNGER 


Presence of cavities in snags retained in forest cutblocks: Do management policies promote 
species retention? Kim T. EVERETT and KEN A. OTTER 


Summer diet of two White-tailed Deer, Odocoileus virginianus, populations living at low and 
high density in southern Quebec CLAUDE DAIGLE, MICHAEL CRETE, LOUIS LESAGE, 
JEAN-PIERRE OQUELLET, and JEAN HUOT 


Winter habitat use by Wolves, Canis lupus, in relation to forest harvesting in west-central Alberta 
GERALD W. KUZYK, JEFF KNETEMAN, and FIONA K. A. SCHMIEGELOW 


Bird communities of the Garry Oak habitat in southwestern British Columbia 
WAYNE R. ERICKSON 


Invertebrate diversity under artificial cover in relation to boreal forest habitat characteristics 
STEPHEN H. FERGUSON and DANIELLE K. A. BERUBE 


Observations of habitat use by Polar Bears, Ursus maritimus, in the Alaskan Beaufort, Chukchi, 
and northern Bering Seas DONALD J. HANSEN 


New records for the Arctic Shrew, Sorex arcticus, and the newly recognized Maritime Shrew, 
Sorex martimensis NEIL D. PERRY, DONALD T. STEWART, ELIZABETH M. MADDEN, 
and THOMAS J. MAIER 


Lichen zonation on coastal rocks on Gwaii Haanas National Park Reserve, Haida Gwaii 
(Queen Charlotte Islands), British Columbia IRWIN M. BRobo and Norm A. SLOAN 


Predaceous water beetles (Coleoptera: Adephaga: Dysticidae, Gyrinidae) collected along the 
Horton and Thelon rivers in the Arctic Central Barrens of Canada 
HELENA V. SHAVERDO and DONNA J. GIBERSON 


Notes 


Collapsing burrow causes death of a Eurasian Beaver, Castor fiber 
Liat R. THOMSEN, FIONA SHARPE, and FRANK ROSELL 


Frequency of tail breakage of the Northern Water Snake, Nerodia sipedon sipedon | KENNETH D. BOWEN 


The Heather Vole, genus Phenacomys, in Alaska 
S. O. MACDONALD, AMY M. RUNCK, and JOSEPH A. COOK 


Extension de Vaire de distribution connue de la Musaraigne fuligineuse, Sorex fumeus, dans le nord-est 
du Quebec JEAN-FRANCOIS DESROCHES and ISABELLE PICARD 
Gulls, Larus spp., foraging at Pink Salmon, Oncorhynchus gobuscha, spawning runs — MAry F. WILLSON 


(continued on inside back cover) 


ISSN 0008-3550 


2004 


368 


376 


The CANADIAN 
FIELD-NATURALIST 


Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada 


re Ave 


Volume 118, Number 4 October—December 2004 


The Ottawa Field-Naturalists’ Club 


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Cover: Lodgepole Pine Dwarf Mistletoe, Arceuthobium americanum, growing on Jack Pine, Pinus banksiana, Growth in Manitoba. 
Photograph by Jacques C. Tardif. See article by Epp and Tardif, pages 595-601. 


MCZ 
LIBRARY 


APR 0 3 2006 


ARVARD 
October-December 2004 


The Canadian Field-Naturalist 


Volume 118, Number 4 


Stranding of a Pygmy Sperm Whale, Kogia breviceps, in the Northern 
Gulf of St. Lawrence, Canada 


LENA Measures!”, BENOIT ROBERGE?, and RICHARD SEARS® 


' Fisheries and Oceans, Maurice Lamontagne Institute, 850 Route de la Mer, Mont-Joli, Québec GSH 324 Canada. (Author 
to whom correspondence should be addressed.) 

2 Parks Canada Agency, Mingan Archipelago National Park Reserve of Canada, 1303 de la Digue, Havre-Saint-Pierre, 
Québec GOG 1P0 Canada 

3 Mingan Island Cetacean Study, Inc., 625, rue du Centre, Longue-Pointe-de-Mingan, Québec GOV 1V0 Canada 


Measures, Lena, Benoit Roberge, and Richard Sears. 2004. Stranding of a Pygmy Sperm Whale, Kogia breviceps, in the 
Northern Gulf of St. Lawrence, Canada. Canadian Field-Naturalist 118(4): 495-498. 


A Pygmy Sperm Whale, Kogia breviceps, stranded alive and later died in the Northern Gulf of St. Lawrence, Quebec, Canada 
on 28 August 2001. This is the northern-most stranding of this species in the western Atlantic. The whale was estimated to be 
approximately 3 m long and a longitudinal section from one tooth (31 mm long and 5.0 mm in diameter) revealed 3.5 growth 


layer groups. ~ 


Key Words: Pygmy Sperm Whale, Kogia breviceps, Gulf of St. Lawrence, Quebec, Canada. 


A Pygmy Sperm Whale (Kogia breviceps) was 
found dead 28 August 2001 on the north shore of the 
Gulf of St. Lawrence. It was observed alive on 27 
August by a local resident, Mme Rose-Annette Blais, 
northwest of St. Charles Island in the Mingan Archi- 
pelago National Park Reserve. The whale was observed 
making slow, undulating movements in shallow water 
close to shore in Trilobites Bay (50°14'07"N; 63°21' 
18"W) and was found dead the next morning on a 
sandy beach in the bay. Personnel from Parks Canada 
were alerted and arrived on site to examine the carcass. 
The whale was estimated to be approximately 3 m long 
(Figure 1). There were various wounds and lacerations 
on the rostrum, flanks and abdomen (Figure 2). Two 
teeth (one broken near the root) collected by Jacques 
A. Thériault and his father, were later submitted for 
analysis. Fisheries and Oceans Canada and the Min- 
gan Island Cetacean Study, Inc. were informed of the 
stranding but before the carcass could be collected 
for necropsy it had been carried off by a rising tide 
and was not seen again. Both teeth were deposited in 
the Maurice Lamontagne Institute collection, Acces- 
sion Number 11629. 

The whale was identified as a Pygmy Sperm Whale 
based on photographs, approximate length (3 m), the 
low, falcate dorsal fin located posterior to the center 
of the dorsum (Figures 1, 2) and size of one intact 
tooth. The tooth was 31 mm long and 5.0 mm in diam- 


eter at its widest point. In the dentine and cementum 
layers there were 3.5 growth layer groups (GLG) 
observed in a longitudinal section of the tooth. It is 
unknown how many GLGs are laid down per year in 
Pygmy Sperm Whales. If similar to the Sperm Whale 
Physeter macrocephalus [1 GLG = 1 year of age (C. 
Lockyer, personal communication; Perrin and Myrick 
1980)], then this animal is almost 4 years old and 
likely an adult (Handley 1966). Ross (1984) reported 
3.5 GLGs in a sexually mature female that stranded 
with a calf. 

The Pygmy Sperm Whale can be confused with 
the Dwarf Sperm Whale (Kogia simus). The latter is 
smaller (2.7 m maximum length) with a taller dolphin- 
like dorsal fin and teeth are less than 30 mm long and 
less than 4.5 mm in diameter (Handley 1966; Leather- 
wood and Reeves 1983). 

Pygmy Sperm Whales are rarely observed in eastern 
Canadian waters and have no COSEWIC (Committee 
on the Status of Endangered Wildlife in Canada) status 
(Baird et al. 1996). Seven, probably eight, Pygmy 
Sperm Whale strandings have been reported prior to 
the present report (Piers 1923; Sergeant et al. 1970: 
Nelson et al. 1991; McAlpine et al. 1997; Lucas and 
Hooker 2000) in Canada over the last century, specif- 
ically in Halifax and on Sable Island in Nova Scotia, 
in Blacks Harbour and Saint-John’s, New Brunswick 
and on the French island of Miquelon near the south- 


495 


496 THE CANADIAN FIELD-NATURALIST Vol. 118 


pau 


FiGure |. Pygmy Sperm Whale beach-cast in Trilobites Bay on 27 August 2001. Length is approximately 3 m. 


west coast of Newfoundland. The present report is the 
northern-most stranding of this species in the western 
Atlantic and the first within the Gulf of St. Lawrence. 

The Pygmy Sperm Whale is a small pelagic, mainly 
deep-water odontocete (occasionally seen motionless 
in surface waters) found widely offshore on continen- 
tal slopes throughout tropical and warm temperate 
waters in the world. Most information on this species 
has been obtained from beach-cast carcasses and re- 
cently from live-stranded and rehabilitated animals in- 
cluding mother-calf pairs or pregnant females (Hiick- 
stadt and Antezana 2001; Scott et al. 2001; Manire et 
al. 2002). Stomach analyses indicate that the Pygmy 
Sperm Whale feeds primarily on cephalopods, crus- 
taceans (shrimp, crab) and fish (Reeves et al. 2002). 
In the western Atlantic most strandings have been 
reported off the southeastern coast of the United States 
(Caldwell and Caldwell 1989; Odell 1991) and live 
individuals have been observed beyond the continental 
shelf and in the Gulf Stream. 

Recently, a rehabilitated Pygmy Sperm Whale 
equipped with a time-depth recorder was released in 
the Gulf Stream off the eastern coast of the United 
States. It was observed for 5 days and remained east 
of the continental shelf break (200 m isobath) but west 
of the eastern edge of the Gulf Stream where sea- 
surface temperatures were 27.8 — 30.0°C (Scott et al. 
2001). Unfortunately, dive depth information was not 


obtained and it is unclear whether the behaviour of 
this individual reflects that of the species in general. 

Odell et al. (1985 as cited in Caldwell and Caldwell 
1989) noted that more strandings of Kogia (K. brevi- 
ceps and K. simus) occur in Florida when the Gulf 
Stream shifts farther offshore. They suggested that indi- 
viduals following prey caught in Gulf Stream eddies 
or rings that suddenly dissipate may become disori- 
ented and, subsequently, strand. Gulf Stream rings 
can occur to the north or south of the Gulf Stream but 
anticyclonic or warm-core rings which are about 
1000 m deep and exist on average 4.5 months, drift 
southwestward to rejoin the Gulf Stream near Cape 
Hatteras (Richardson 1976; Wiebe 1976). Gulf stream 
rings have unique physical, chemical and biological 
characteristics (Wiebe 1976; Joyce and Wiebe 1983; 
Craddock et al.1992). Although warm-core rings do 
not ride onto shallow continental shelves, they can 
push slope water onto the shelf or entrain shelf water 
into slope waters. Such shelf water entrainment was 
suggested as providing suitable cephalopod habitat 
which attracted Sperm Whales in the vicinity of a 
warm-core ring off Georges Bank (Griffin 1999). 

The occurrence of a Pygmy Sperm Whale in the 
Gulf of St. Lawrence, especially in the northern Gulf, 
seems unusual as water temperatures are relatively cold 
[sea-surface temperatures are generally 12 — 18°C in 
August (Vigeant 1987*)]. The minimum temperature 


2004 


MEASURES, ROBERGE, AND SEARS: STRANDING OF PYGMY SPERM WHALE 


497 


FiGuRE 2. Pygmy Sperm Whale in shallow water in Trilobites Bay (same specimen as in Figure 1). Note relative position of 
dorsal fin on trunk. 


tolerated by Pygmy Sperm Whales is unknown. Annu- 
al mean air temperatures in most of the northwest At- 
lantic, especially in the southern Gulf of St. Lawrence, 
and sea-surface water temperatures throughout eastern 
Canadian waters in 2001 were generally warmer than 
normal (Drinkwater et al. 2002a*). In some areas, the 
upper 30 m in the central and southern Gulf of St. 
Lawrence show monthly anomalies of 3 - 4°C (Drink- 
water et al. 2002b*; Gilbert 2002*). The Pygmy Sperm 
Whale may have swam and drifted northeast follow- 
ing prey in the Gulf Stream as suggested by Fraser 
(1974), perhaps using warm-core rings as “stepping- 
stones” (Peter Wiebe, personal communication) or 
thermal fronts associated with warm-core rings as 
observed by Griffin (1999). It may have entered the 
Gulf of St. Lawrence in warm surface waters on the 
Cape Breton side of the Cabot Strait when the Cape 
Breton Current slows in summer (El-Sabh 1977). Dur- 
ing a DFO research cruise in the Gulf of St. Law- 
rence in August 2001 squid (/lex illecebrosus) were 
collected but numbers of specimens collected were 
not considered exceptional (D. Archambault, DFO, 
unpublished data). 


Acknowledgments 

We thank R-A. Blais and J. A. Thériault for assis- 
tance in this stranding and providing teeth. We also 
thank S. Plamondon of Parks Canada who provided 
photographs and J.-G. Gosselin of Fisheries and Oceans 
Canada who determined the number of GLGs by cut- 
ting the tooth. We appreciate that P. Best and J. Mead 
examined the photographs and concurred with our 
identification of the species. 


Documents Cited (marked * in text) 

Drinkwater, K. F., R. G. Pettipas, and W. M. Petri. 2002a. 
An overview of meteorological, sea ice and sea-surface 
temperature conditions off Eastern Canada during 2001. 
Canadian Science Advisory Secretariat Research Docu- 
ment 2002/048. 

Drinkwater, K. F., R. G. Pettipas, and W. M. Petri. 2002b. 
Physical environmental conditions in the Southern Gulf 
of St. Lawrence during 2001. Canadian Science Advisory 
Secretariat Research Document 2002/047. 

Gilbert, D. 2002. Oceanographic conditions in the Gulf of 
St. Lawrence in 2001 : physical oceanography. DFO Sci- 
ence Stock Status Report. 


498 


Vigeant, G. 1987. Température mensuelle de ]’eau en surface 
dans l’estuaire et le Golfe du Saint-Laurent. Service de 
1’ Environnement Canada. 


Literature Cited 

Baird, R. W., D. Nelson, J. Lien, and D. W. Nagorsen. 
1996. The status of the pygmy sperm whale, Kogia brevi- 
ceps, in Canada. Canadian Field-Naturalist 110: 525-532. 

Caldwell, D. K., and M. C. Caldwell. 1989. Pygmy sperm 
whale Kogia breviceps (de Blainville, 1838): dwarf sperm 
whale Kogia simus Owen, 1866. pages 235 — 260 in Hand- 
book of marine mammals. Edited by S. H. Ridgway and 
R. Harrison. Academic Press, London. 

Craddock, J. E., R. H. Backus, and M. A. Daher. 1992. 
Vertical distribution and species composition of midwater 
fishes in warm-core Gulf Stream meander/ring 82-H. Deep 
Sea Research 39: S203-S218. 

El-Sabh, M. I. 1977. Oceanographic features, currents, and 
transport in Cabot Strait. Journal of the Fisheries Research 
Board of Canada 34: 516-528. 

Fraser, F. C. 1974. Report on Cetacea stranded on the British 
coasts from 1948 to 1966. British Museum (Natural History) 
14. 

Griffin, R. B. 1999. Sperm whale distributions and commu- 
nity ecology associated with a warm-core ring off Georges 
Bank. Marine Mammal Science 15: 33-51. 

Handley, C. O., Jr. 1966. A synopsis of the genus Kogia 
(pygmy sperm whales), pages 62 — 69 in Whales, dol- 
phins and porpoises. Edited by K. S. Norris. University of 
California Press, Berkeley and Los Angeles, Calif. 

Hiickstaédt, L., and T. Antezana. 2001. An observation of 
parturition in a stranded Kogia breviceps. Marine Mammal 
Science 17: 362-365. 

Joyce, T., and P. Wiebe. 1983. Warm-core rings of the Gulf 
Stream. Oceanus 26: 34-44. 

Leatherwood, S., and R. R. Reeves. 1983. The Sierra Club 
handbook of whales and dolphins. Sierra Club Books, San 
Francisco. 

Lucas, Z. N., and S. H. Hooker. 2000. Cetacean strandings 
on Sable Island, Nova Scotia, 1970 — 1998. Canadian Field- 
Naturalist 114: 45-61. 

Manire, C. A., L. Byrd, H. L. Rhinehart, P. Cunningham- 
Smith, and D. R. Smith. 2002. Subacute atropine toxicity 
in a pygmy sperm whale, Kogia breviceps. Journal of Zoo 
and Wildlife Management 33: 66-72. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


McAlpine, D. F., L. D. Murison, and E. P. Hoberg. 1997. 
New records for the pygmy sperm whale, Kogia breviceps 
(Physeteridae) from Atlantic Canada with notes on diet 
and parasites. Marine Mammal Science 13: 701—704. 

Nelson, D., A. Desbrosse, J. Lien, P. Ostromm, and R. 
Seton. 1991. A new stranding record of the pygmy sperm 
whale, Kogia breviceps, in waters off Eastern Canada. 
Canadian Field-Naturalist 105: 407-408. 

Odell, D. K. 1991. A review of the Southeastern United States 
Marine Mammal Stranding Network: 1978 — 1987. pages 
19 — 23 in Marine Mammal Strandings in the United States. 
Edited by J. E. Reynolds II and D. K. Odell. NOAA 
Technical Report NMFS 98. 

Perrin, W. F., and A. C. Myrick, Jr. Editors. 1980. Growth 
of odontocetes and sirenians: problems in age determina- 
tion. Reports of the International Whaling Commission 
Special Issue 3: 1-229. 

Piers, H. 1923. Accidental occurrence of the pygmy sperm 
whale (Kogia breviceps) on the coast of Nova Scotia; an 
extension of its known range, with remarks on the proba- 
bility of the former presence in these waters of the true 
sperm whale (Physeter macrocephalus). Proceedings and 
Transactions of the Nova Scotia Institute of Science 15: 
95-114. 

Reeves, R. R., B. S. Stewart, P. J. Clapham, and J. A. 
Powell. 2002. National Audubon Society guide to marine 
mammals of the world. Alfred A. Knopf, New York. 

Richardson, P. 1976. Gulf Stream rings. Oceanus 19: 65-68. 

Ross, G. J. B. 1994. The smaller cetaceans of the south east 
coast of southern Africa. Annals of the Cape Provincial 
Museums (Natural History) 15: 173-410. 

Scott, M. D., A. A. Hohn, A. J. Westgate, J. R. Nicolas, B. 
R. Whitaker, and W. B. Campbell. 2001. A note on the 
release and tracking of a rehabilitated pygmy sperm whale 
(Kogia breviceps). Journal of Cetacean Research Man- 
agement 3: 87- 4. 

Sergeant, D. E., A. W. Mansfield, and B. Beck. 1970. 
Inshore records of cetacea for eastern Canada, 1949 — 68. 
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1903-1915. 

Wiebe, P. 1976. The biology of cold-core rings. Oceanus 
19: 69-76. 


Received 14 July 2003 
Accepted 15 July 2004 


Seed Dispersal by Brown Bears, Ursus arctos, in Southeastern Alaska 


Mary FE. WILLSON! and Scott M. GENDE? 


'5230 Terrace Place, Juneau, Alaska 99801 USA. 
National Park Service, Glacier Bay Field Station, 3100 National Park Road, Juneau, Alaska 99801 USA 


Willson, Mary F., and Scott M. Gende. 2004. Seed dispersal by Brown Bears, Ursus arctos, in southeastern Alaska. Canadian 
Field-Naturalist 118(4): 499-503. 


Mammals often consume fleshy fruits and disperse significant quantities of the enclosed seeds. In southeastern Alaska, 
Brown Bears (Ursus arctos) are among the most important dispersers of seeds for the numerous plant species producing 
fleshy fruits, because these bears are abundant, often eat large quantities of fruit, and commonly excrete seeds in germinable 
condition. Scat analyses showed that Brown Bears on Chichagof Island ate increasing quantities of fruit through summer 
and fall. Scats commonly contained several thousand seeds, often of two or more species. Four kinds of seeds of fleshy- 
fruited plants that normally grow in forest understory germinated at similar levels when experimentally deposited (in bear 
scats) in the two most common habitats (forest and muskeg), suggesting that habitat distribution of these plants is not deter- 
mined simply by germination patterns. Although seed passage through bear digestive tracts and the composition of scats are 
known to affect germination rates to some degree, the most important role of bears in seed dispersal is probably transport. 


Key Words: Brown Bears, Ursus arctos, Southeastern Alaska, seed dispersal, fleshy fruits, Rubus, Ribes, Oplopanax, Vaccinium, 


Streptopus. 


Coastal rainforest ecosystems in southeastern Alaska 
include many shrubs and herbs with fleshy fruits, 
commonly consumed by vertebrates that disperse the 
enclosed seeds (Willson 1991; Traveset et al. 2004). 
Potential dispersal agents include corvids, thrushes, 
warblers, waxwings, Pine Marten (Hickey et al. 1999) 
and bears (Willson 1993, 1994). Bears are important 
dispersers of seeds, because they can consume large 
quantities of fruit (Welch et al. 1997; Farley and Rob- 
bins 1995) and often range over considerable dis- 
tances (Patten 1993), distributing seeds far from the 
source. Furthermore, bears are numerous in south- 
eastern Alaska, with density reaching several hundred 
bears per 1000 km — perhaps as many as 700 or even 
more in some areas (Miller et al. 1997; J. Whitman, 
personal communication). The abundance of Brown 
Bears (Ursus arctos) in southeastern Alaska provides 
an opportunity to examine the consequences of bear 
frugivory for seed dispersal. 

The purpose of this paper is to describe the phenol- 
ogy of Brown Bear frugivory, patterns of scat deposition 
on a small spatial scale, seed abundance and composi- 
tion in bear scats, and field tests of seed germination 
from bear scats. We then integrate this information 
with previous reports, in order to describe what is 
known about this ecologically important interaction 
in southeastern Alaska. 


Study Area and Methods 

Field work was conducted on northern Chichagof 
Island in the Alexander Archipelago, at the southern end 
of Port Frederick (approximately 58°N, 135°30'W). 
The only bears on this island are Brown Bears. The 
rainforest is composed chiefly of Sitka Spruce (Picea 


sitchensis) and hemlock (Tsuga spp.), with occasional 
muskegs (bogs) of Sphagnum moss, scattered Lodge- 
pole Pines (Pinus contorta), and low shrubs. Brown 
Bear scats were collected from abandoned logging 
roads, at less than 300 m elevation, in the Neka River 
valley and near Salt Lake Bay on opposite sides of 
Port Frederick. During each road survey, all scats were 
cleared from the road, so that the subsequent survey 
recorded only scats deposited in the between-survey 
interval. The roads passed through forests of differing 
ages, clearcuts, and muskegs. 


Seed composition of scats 

The frequency of occurrence of seeds, and the prin- 
cipal kinds of seeds present, were recorded in June- 
September 1990-1993 in 4378 scats. A subset of 570 
seed-containing scats was collected in August and 
September 1990 and 1991, for quantitative estimates 
of seed abundance. Each of these scats was mixed to 
distribute the seeds throughout, and a subsample aver- 
aging 24% + 2% (S. E.) of the total scat mass was 
taken. In the weighed subsample, the seeds were iden- 
tified to genus and counted; the number of seeds in 
the entire scat was then estimated by extrapolation. 
The frequency of seeds in scats through the summer 
and early fall was examined, using | June as Day 1. 
June scat collections represent the previous spring 
months; collections in July, August, and September 
represent only the preceding 30 days, approximately. 


Seed germination 

In order to compare seed germination in two prin- 
cipal habitats, over 150 of the scats for which total 
seed content was estimated were placed in two common 
habitats (muskeg or forest). Scats were “planted” in 


499 


500 


transects in each habitat, in whatever microhabitats 
were available at about 2 m intervals. Muskeg and for- 
est transects near Salt Lake Bay were established in 
late summer, 1990, and in the Neka Valley in 1991. 
Seedlings were counted in the planted scats annually 
(in August) for three (Neka) or four (Salt Lake Bay) 
years. For seedling censuses, each planted scat was 
covered with a grid, and seedlings were counted in a 
random selection of the grid squares. The proportion 
of grid squares sampled usually ranged from 20% to 
about 50% of the total scat. Emerging seedlings were 
counted individually when possible, but so many seeds 
germinated in some squares that it was impossible to 
count them without destructive sampling, in which case 
the abundance of seedlings was estimated visually. 
The total number of emerging seedlings per scat was 
extrapolated from this subsample. Curious bears and 
heavy fall rains disturbed some of the planted scats and 
thus reduced the usable sample size slightly. Because 
it was impossible to mark individual seedlings in the 
very dense clusters, the data are simply estimates of 
annual germination, not of survivorship of seedlings. 
Mortality of seedlings was very high and only a few 
survived more than one year. We did not observe rodent 
and bird predation on seeds in the experimental tran- 
sects, although seed predation was sometimes evident 
in scats on the road system. 

The estimated percent of seeds germinating each 
year was compared between habitats using a repeated- 
measures ANOVA on arc-sine transformed data (PROC 
GLM, SAS Systems Inc., Cary, North Carolina). 
Because the power of this test was low (two sites, 
two habitats), our results were re-examined, to see if 
the outcome was merely a result of the few degrees 
of freedom in the ANOVA. For this we used a t-test 
(in which the planted scats were the replicates, so there 
were more degrees of freedom in the test) to test for 
differences in seedling germination between habitats 
for the first and the last years at each site (no seedlings 
in the censused grid squares survived for the entire 
length of the experiment). 


Results 
Phenology of fruit consumption 

Scats collected in early summer contained mostly 
vegetable fiber and occasional deer remains (hair, bone, 
hoof) with few seeds, as expected, because few fruits 
are available at that time of year (some overwintered 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


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Ficure |. Seasonal changes, for four years, in the percent of 
bear scats containing seeds, Chichagof Island, south- 
eastern Alaska. R, = 0.79, p < 0.001. 


Viburnum and Maianthemum; some early Rubus spec- 
tabilis). The frequency of seed-containing scats in- 
creased through the summer, from less than 1% in 
mid-June to over 90% in late August and early Sep- 
tember (R, = 0.79, p < 0.001, n = 12; Figure 1). 


Scat distribution at a small spatial scale 

In addition to widespread deposition of scats on the 
landscape (Patten 1993), defecations by a moving bear 
can spread multiple deposits of a single defecation over 
several meters, which could be ecologically important. 
A survey of 1163 “scat trails” revealed that 42% were 
multiple deposits, with 3% of them comprised of over 
20 separate droppings spread over as much as 84 m. 


Seed composition in scats 

Seeds were identified at least to genus; in some 
cases, the genus was represented by only one species 
in this area (Sambucus racemosa, Rubus spectabilis, 
Oplopanax horridum). In the case of Ribes, the species 
was most likely bracteosum, which grew commonly 
in this area. Likewise, Streptopus amplexifolius was 
far more common in this area than two congeners. 
Both Vaccinium ovalifolium and V. alaskense were 
present and were not distinguishable in the scats. The 
most commonly occurring seeds were Oplopanax, 
Vaccinium, Ribes, Rubus, and Streptopus (Table 1). 

Seeds had a high probability of being deposited 
not only with numerous conspecifics but also with at 


TABLE 1. Summary of frequency of occurrence of common kinds of seeds in scats on northern Chichagof Island. Data are 
the number of months that the frequency of occurrence equaled or exceeded the stated value. 


Kind of seed 
Vaccinium Ribes Oplopanax Rubus Streptopus 
Number of months = 20% 5 5) 1 1 
3 ia “8 


Number of months = 50% 3} 


2004 


WILLSON and GENDE: SEED DISPERSAL BY BROWN BEARS 


TABLE 2. Average (S.E.) number of seeds per scat on northern Chichagof Island. N = 570 scats. 


Year Month 
Vaccinium Ribes 
1990 July 3581 (667) 914 (767) 
August 4655 (461) 700 (165) 
September 2534 (468) 1918 (281) 
199] July 4375 11(933) 2.02817 (-) 
August 7193 (760) 364 (109) 
September 4329 (795) 1302 (293) 
4445 1971 
x number of seeds/fruit* 47 12 
x number of fruits 95 89 


*from Traveset et al. 2004. 


least one other kind of seed: in monthly samples, a 
median of 60% (range 16-87%) of scats contained at 
least two kinds of seeds, and sometimes as many as 
five kinds. Vaccinium and Ribes were consistently the 
most abundant seeds in scats collected (Table 2). In 
most months, Rubus was next most abundant, followed 
by Oplopanax and others. The maximum numbers 
per scat for each of the genera listed in Table 2 were 
approximately 37 000 Vaccinium, 14 500 Ribes, 7600 
Rubus, 2000 Oplopanax, 1900 Sambucus, and 400 
Streptopus. In addition to those represented in Table 2, 
other kinds of seeds appeared in the scats sporadical- 
ly (from fleshy fruits, Viburnum, Maianthemum; non- 
fleshy fruits, Kalmia, Heracleum, Carex, grass). 
Because the number of seeds per fruit differs greatly 
among genera (Traveset et al. 2004), the approximate 
number of consumed fruits of each type per scat differs 
markedly from the abundances of seeds (Table 2). 
The average scat represented only approximately 89 
— 116 fruits of the most commonly consumed types. 


Seed germination 

Germination patterns were examined only for Vac- 
cinium, Ribes, Rubus, and Oplopanax. A few Vaccinium 
seeds germinated in the year they were planted (<1% 
of all seeds estimated to be present); germination levels 
were generally low (<6%) over the next four years 
(Table 3). Annual germination of Oplopanax was also 
rather low (<18%). In contrast, both Ribes and Rubus 
germinated relatively well (up to 31-34% per year), 
with the highest levels of germination often in the 
second year after planting. All these kinds of seeds were 
clearly capable of dormancy for at least four years. 

There were no differences in germination percent- 
ages between habitats for any species (ANOVA; range 
of p-values: 0.17 — 0.82). The subsequent exploratory 
t-test also revealed no differences between habitats 
except for Vaccinium in one site — one statistically sig- 
nificant outcome in 16 tests, which might happen by 
chance alone. Variation among scats planted within a 
habitat was relatively large, but it probably reflected the 
biological reality of variation on a small spatial scale. 


501 
Kind of seed 
Oplopanax Rubus Streptopus Sambucus 
325). 1G18) 383 (91) = = 
134 (22) 123) (182) 307 (8) 126 (42) 
360 = (42) 830 (291) 41 (15) 88 (33) 
3K (25) 650 (137) 47 (26) - 
228 (40) 583 (120) 81 (20) - 
374 (76) 54 = (20) 56 (12) 683 (606) 
231 634 5] 110 
2 42 22 3 
116 15 2 37 
Discussion 


Bears are well-known frugivores, consuming many 
species of fleshy fruits (Auger et al. 2002; Willson 
1993). Fruits can be an important resource for prepar- 
ing for winter dormancy and lactation (females give 
birth in winter dens), and berry-crop failures can lead 
to lower reproductive success (Rogers 1987). Bears can 
consume many kilograms of fruits in a day (Welch et 
al. 1997; Farley and Robbins 1995). Although a single 
bear scat in the present study often contained thou- 
sands of seeds, this represented relatively few fruits 
and therefore only a small proportion of possible 
daily consumption. 

Composition of the fecal deposit can affect seed 
germination and seedling growth (Traveset et al. 2001), 
and the concentration of seeds in scats can lead to post- 
dispersal seed predation and secondary dispersal via 
scattering (Bermejo et al. 1998). Germination of seeds 
passed by captive bears was generally similar to or 
faster than that of seeds extracted from fruits by 
researchers, although germination of some species 
was quite low in all conditions and was sometimes 
dependent on the substrates on which the seeds hap- 
pen to land (Traveset and Willson 1997, 1998). But 
rapid germination is not necessarily beneficial, if seed 
dormancy is indeed an adaptation for dispersal in time. 

The most important consistent effects of frugivory 
by bears on seed dispersal may be transport, rather 
than shifts of germination behavior of seeds passed 
through the digestive tract. In a companion study, the 
probability of seed deposition by Brown Bears on 
Chichagof Island was greatest at 600-900m from a 
source (Patten 1993). The greatest distance estimated 
for seed transport by Brown Bears was over three km 
(Patten 1993), and even though few seeds are likely 
to be carried this far from their source, the extended 
tail of the seed distribution pattern can be dispropor- 
tionately important for dispersal ecology (Portnoy and 
Willson 1992) and plant community ecology (Vellend 
et al. 2003). Little habitat preference was shown by the 
bears tracked on Chichagof, except for an avoidance 
of bare-rock areas, so the probability of seed deposi- 


502 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE 3. Estimated average percent of seeds germinating per year in two common habitats at two sites on Chichagof Island, 


southeastern Alaska. 


Site Habitat Year Kind of seed 
Vaccinium Ribes Oplopanax Rubus 
Neka Forest 1992 3.9 (0.9) 0.2 (0.2) <0.1 = 22.5 (7.2) 
n= 30 1993 2.8 (0.5) 30:8. e@li5e7) NS:42) HGLOSI) 28.5 (9.8) 
1994 313 (0.9) 14.7 (9.0) 9.5 (4.9) 13.6 (4.8) 
Muskeg 1992 5.9 (2:2) - Lal (1.0) 20.7 Ga) 
n= 30 1993 2.9 (1.0) - 10.2 (8.4) 19.7 GS) 
1994 1.2 (0.4) — TIEOe., .LO%6) 5.6 68) 
Salt Lk Bay Forest 1991 2.4 (0.9) 19 x CluES)) 55} A) EOF (933) 
n=60 1992 3.6 (0.8) 30.8 (16.2) 1.6 (0.8) 34.2 (8.7) 
1993 1.8 (0.3) DES (es) iNet (0.5) 19.7 (6.9) 
1994 ES (0.2) 1.0 (0.6) 3.0 (1.3) 9.8 (4.8) 
Muskeg 1991 1.1 (0.5) 0.5 (0.5) 4.6 (2.7) heal (5.1) 
n=49 1992 3.0 (1.0) 25.6 (10.4) 122, (0.7) 19.6 (6.9) 
1993 1.8 (0.7) 6.3 (52) 0.9 (0.7) 10.7 (4.5) 
1994 1.6 (0.8) 0.3 (0.2) 0.6 (0.4) 4.3 (3.0) 


tion was proportional to habitat availability, including 
logged areas (Patten 1993). Because forest was the 
common habitat, this meant that the forest fruits, such 
as most of the species examined here, had a fairly high 
probability of deposition in the appropriate habitat 
(Patten 1993). On a smaller scale, defecation by mov- 
ing bears could increase the probability that seeds from 
one defecation episode are deposited in more than one 
habitat or microhabitat, as well as decrease the inten- 
sity of seedling competition. 

Germination of several forest and forest-edge species 
was similar in forest and muskeg, indicating that habi- 
tat distribution of these shrubs is not determined by 
germination requirements at the habitat scale. The large 
variances of the estimates of germination level not 
only decreased the probability of detecting habitat 
differences, but they also suggest that microhabitats 
of seed deposition may be more important in deter- 
mining germination success than habitats. Mortality 
of seedlings was very high, in part an apparent result 
of desiccation during periods of warm, sunny weather. 
Especially at high densities, intra- and interspecific 
competition was undoubtedly intense and probably 
contributed to seedling mortality. However, occasion- 
ally small groups of seedlings persisted and were still 
alive when field work ended; similar “bear gardens” 
were also found in natural settings (Willson 1994). 

Bears are good seed dispersers: they commonly 
transport germinable seeds up to hundreds of meters 
from a source, often in appropriate habitat, simulta- 
neously with a load of fertilizer that may influence 
germination and growth. Although scats containing 
numerous seeds produce conditions of high seedling 
competition, several factors tend to reduce the impact 
of competition on the effectiveness of dispersal. Ger- 
mination may be spread over several years. Relatively 
rapid passage through the gut and multiple defecations 
per day within a bear’s home range spread scats over 


the landscape. Scat trails of moving bears spread out 
the seeds on a more local scale. Bears may be more 
reliable dispersal agents for many kinds of seeds than 
most frugivorous birds in this area, since the coastal 
region is not a major flyway for migratory forest birds. 


Acknowledgments. 

This project was funded by the Pacific Northwest 
Research Station, U.S. Forest Service. We are grate- 
ful to J. Zasada for engaging us with bear-fruit inter- 
actions, to our numerous field assistants, and to A. 
Traveset for comments on the ms. 


Literature Cited 

Auger, J., S. E. Meyer, and H. L. Black. 2002. Are American 
black bears (Ursus americanus) legitimate seed dispersers 
for fleshy-fruited shrubs? American Midland Naturalist 
147: 352-367. 

Bermejo, T., A. Traveset, and M. F. Willson. 1998. Post- 
dispersal seed predation in the temperate rainforest of 
Southeastern Alaska. Canadian Field-Naturalist 112: 510- 
512. 

Farley, S. D., and C. T. Robbins. 1995. Lactation, hiberna- 
tion, and mass dynamics of American black and grizzly 
bears. Canadian Journal of Zoology 73: 2216-2222. 

Hickey, J. R., R. W. Flynn, S. W. Buskirk, K. G. Gerow, 
and M. F. Willson. 1999. An evaluation of a mammalian 
predator, Martes americana, as a disperser of seeds. Oikos 
87: 499-508. 

Miller, S. D., G. C. White, R. A. Sellers, H. V. Reynolds, 
J. W. Schoen, K. Titus, V. G. Barnes Jr., R. B. Smith, 
R. R. Nelson, W. B. Ballard, and C. C. Schwartz. 1997. 
Brown and black bear density estimation in Alaska using 
radiotelemetry and replicated mark-resight techniques. 
Wildlife Monographs 133: 1-55. 

Patten, L. A. 1993. Seed dispersal patterns generated by brown 
bears (Ursus arctos) in southeastern Alaska. MS thesis, 
Washington State University, Pullman, Washington. 

Portnoy, S., and M. F. Willson. 1992. Seed dispersal curves: 
the behavior of the tail of the distribution. Evolutionary 
Ecology 7: 25-44. 


2004 


Rogers, L. L. 1987. Effects of food supply and kinship on 
social behavior, movements, and population growth of 
black bears in northeastern Minnesota. Wildlife Mono- 
graphs 97: 1-72. 

Traveset, A., and M. F. Willson. 1997. Effect of birds and 
bears on seed germination of fleshy-fruited plants in tem- 
perate rainforests of southeastern Alaska. Oikos 80: 89-95. 

Traveset, A., and M. F. Willson. 1998. Ecology of the fruit- 
colour polymorphism in Rubus spectabilis. Evolutionary 
Ecology 12: 331-345. 

Traveset, A., T. Bermejo, and M. F. Willson. 2001. Effect of 
manure composition on seedling emergence and growth of 
two common shrub species of southeastern Alaska. Plant 
Ecology 155: 29-34. 

Traveset, A., M. F. Willson, and M. Verdi. 2004. Charac- 
teristics of fleshy fruits in southeastern Alaska: Phyloge- 
netic comparison with fruits from Illinois. Ecography 27: 
41-48. 


WILLSON and GENDE: SEED DISPERSAL BY BROWN BEARS 


Vellend, M., J. A. Myers, S. Gardescu, and P. L. Marks. 
2003. Dispersal of Trillium seeds by deer: implications for 
long-distance migration of forest herbs. Ecology 84: 1067- 
1072. 

Welch, C. A., J. Keay, K. C. Kendall, and C. T. Robbins. 
1997. Constraints on frugivory by bears. Ecology 78: 1105- 
1119. 

Willson, M. EF. 1991. Dispersal of seeds by frugivorous animals 
in temperate forests. Revista Chilena de Historia Natural 
64: 537-554. 

Willson, M. F. 1993. Mammals as seed-dispersal mutualists 
in North America. Oikos 67: 159-176. 

Willson, M. F. 1994. Bear gardens. Pacific Discovery 47(1): 
8-14. 


Received 15 July 2003 
Accepted 23 October 2004 


A Mapping of the Present and Past Forest-types of Prince Edward Island 


D. G. SoBey! AND W. M. GLEN? 


! School of Applied Biological and Chemical Sciences, University of Ulster, Jordanstown, Northern Ireland BT37 OQB United 
Kingdom. (Research Associate of the Institute of Island Studies, University of Prince Edward Island, Charlottetown, 
Prince Edward Island) 

2 Natural Resources Division, Department of Agriculture and Forestry, PO Box 2000, Charlottetown, Prince Edward Island 
C1A 7N8 Canada 


Sobey, D. G., and W. M. Glen. 2004. A mapping of the present and past forest-types of Prince Edward Island. Canadian 
Field-Naturalist 118(4): 504-520. 


Our aim was to produce maps showing the distribution on Prince Edward Island of five forest-types previously identified 
from a TWINSPAN analysis of ground flora data collected at 1200 sampling points in a field survey. For this purpose we 
had available two databases: one on the composition of the tree canopy of 82 957 forest stands, as determined by photo- 
interpretation of a 1990 aerial, photographic survey of the island; the other on the drainage properties of the same stands 
from a published soil survey. The tree canopy and drainage criteria for sorting these stands into five stand-types were chosen 
in the light of the equivalent properties of the TWINSPAN forest-types as evident from the field survey. These criteria were 
perfected in four trial computer-sortings, followed by the computer-printing of maps showing the distribution of the stand- 
types. These maps, which were then evaluated by comparing them with the properties of the TWINSPAN forest-types, are the 
first fine-scale maps of the main forest-types of the island. They reveal that, of the three “primary” forest-types, the upland 
hardwood forest occurs especially in the central and south-eastern hill-lands, as well as in scattered parcels elsewhere, 
whereas the Black Spruce forest and the wet species-rich woodland occur primarily in areas of lower elevation in the east 
and west of the island. The two forest-types resulting from human disturbance, the White Spruce woods and the “disturbed 
forest”, have a more scattered distribution, with the White Spruce woods being found especially in the central and eastern 
parts of the island and the disturbed forest in the west and east of the island. A secondary aim was to map the conjectured 
distribution before European settlement of the three primary forest-types: two maps have been produced, one showing the 


distribution of upland hardwood forest, the other of the wet forest-types. 


Key Words: forest classification, forest mapping, forest history, Prince Edward Island. 


Between 1990 and 1992 the Prince Edward Island 
Forestry Division carried out a comprehensive Forest 
Biomass Inventory of the province, a component of 
which was a field survey in 1991 at 240 randomly- 
selected forest sites. At each of the sites the percentage 
cover of the ground flora species in 4-m? plots was 
recorded at five sampling points. The availability of 
such ground flora data from such a large number of 
plots (1200 in total) presented an unprecedented oppor- 
tunity for the study and analysis of the woodland 
ground vegetation and forest-types of Prince Edward 
Island. 

An earlier paper (Sobey and Glen 2002) presented 
an analysis of the ground flora data in 1127 of the 
plots using two multivariate techniques: TWINSPAN 
— two-way indicator species analysis (a classification 
technique), and DECORANA -— detrended correspon- 
dence analysis (an ordination technique). TWINSPAN 
led to the recognition of five forest-types, each char- 
acterized by particular tree species and soil drainage 
properties: (1) a wet species-rich swamp-type wood- 
land, (2) upland hardwood forest, (3) Black Spruce 
(Picea mariana) forest, (4) old field White Spruce 
(Picea glauca) woods, and (5) disturbed conifer-dom- 
inated forest. It was conjectured that the first three of 
these forest-types were heavily modified descendants 


of forest-types occurring at the time of European set- 
tlement, while the two latter appeared to be largely the 
product of successional processes associated with the 
effects of human disturbance and forest clearance. 
The DECORANA ordination provided further sup- 
port for the overall importance of soil drainage and 
human disturbance as the principal factors responsi- 
ble for the differences between the five forest-types. 
The aim of this paper is to expand the results of the 
plant community analysis beyond the 1200 sampling 
points to include the whole forested area of the island, 
in the form of maps showing the total distribution of the 
five forest-types on the island. Since the forest-types 
were initially segregated and defined by TWINSPAN 
on the basis of the composition of their ground flora, 
the ideal mapping approach would have been to use 
the ground flora composition of all forest stands on the 
island as the basis for mapping the distribution of the 
forest-types. However, such data are not available for 
the whole forested area of the island and are never 
likely to be. The forest-types thus had to be mapped 
using data that showed a correlation with the ground 
flora composition (1.e., data on the composition of 
the tree canopy) and also for some, data on soil 
drainage properties. It is fortunate that there were avail- 
able two comprehensive relevant databases for the 


504 


2004 


whole island: the more important was a database on the 
tree species composition of all forest stands on the 
island, based on an aerial photographic survey carried 
out in mid-summer 1990; the other was the Prince 
Edward Island Soil Survey (MacDougall et al. 1988) 
which classified the soils of the island into 44 soil series 
and mapped their distribution (at a scale of 1:10 000) 
over the whole island. The soil survey proved useful in 
the forest mapping because some of the TWINSPAN 
forest-types had high levels of association with partic- 
ular soil drainage properties. 

It should be noted that none of the previous attempts 
at forest classification and mapping that include Prince 
Edward Island (Stilgenbauer 1929; Halliday 1937; 
Rowe 1959; Loucks 1962) aimed to map what was 
actually present on specific sites, nor do they contain a 
high level of detail or accuracy. Halliday’s and Rowe’s 
maps, being part of national studies, show no internal 
differentiation at all in the forests of the island. Loucks 
(working at the level of Maritime forests), did sub- 
divide the island’s forests: he placed them in three 
“forest districts” but the boundaries of these are very 
generalized on his map. One comprised the hardwood 
areas of the central and eastern parts of the island; the 
other two were coniferous districts: one in the west 
along the shores of Northumberland Strait, the other 
running along the length of the northern shore. The 
map showing the greatest detail (Stilgenbauer 1929) 
is actually the earliest of the four, but Stilgenbauer’s 
descriptions of his “forest belts” are brief and quali- 
tative and the criteria he used for delimiting them are 
not given. In addition to these mapping studies, there 
is a useful descriptive summary of the forest-types of 
the island in Erskine (1960), and in Appendix 1 the 
stand-types emerging from this study are equated with 
Erskine’s descriptions. 

The principal objective of this study was thus, mak- 
ing use of two comprehensive databases, to map the 
total distribution on Prince Edward Island of the five 
forest-types recognized in a TWINSPAN analysis, and 
to further verify the validity of the maps by comparing 
the properties of the mapped stands with those of the 
TWINSPAN forest-types. Such a mapping will not only 
give us a clear picture of the current actual distribution 
on the ground of the main forest-types on the island, 
but should also assist us in understanding the factors 
responsible for their distributions. A secondary objective 
(on the basis of any stand and soil relationships that 
might emerge) was to attempt to extend the mapping 
back into historical time in order to obtain a picture of 
the possible distribution of the forest-types before the 
advent of the large scale forest clearance that began 
in the eighteenth century with the beginning of Euro- 
pean settlement on the island. 


Methods 
The methodology leading to the description of the 
forest-types and their mapping makes use of several 


SOBEY AND GLEN: PRESENT AND PAST FOREST-TYPES OF P.E.I. 


large independently obtained databases and involves 
a number of different stages in processing and analysing 
the data they contain. Figure | shows these databases 
and stages in diagrammatic form. The end-product, 
computer-produced maps of the five forest-types, is 
the result of a computer-sorting of 82 957 demarcated 
stands of “high forest” (i.e., excluding alder woods, 
plantations, clear-cut and burned areas) into five forest- 
types on specific tree canopy and soil drainage criteria. 
In the description of the various steps in the analysis, the 
text that follows is linked to the stages shown in Fig- 
ure | by Roman numerals in square brackets, e.g.: [I]. 


The databases: 
1. The 1991 field survey: ground flora and tree canopy 
data 

Of the vast amount of data collected at the 1200 sam- 
pling points in the 1991 field survey, two data sets, 
one on the ground flora [II], the other on the tree can- 
opy [III], are of relevance to this report. We present 
here only a summary of the methods used to collect 
these data — full field methods are given in Sobey and 
Glen (2002). 

The data were collected between June and Sep- 
tember 1991 at 1200 ground flora plots located at 
240 randomly-selected forest sites. At each sampling 
point the tree canopy was assessed using a “point 
sampling” technique involving the use of a “variable- 
radius plot” centred on the sampling point (Watts 
1983). From these field data, the percentage contribu- 
tion of each tree species to the total woody biomass (to 
the nearest 10%) in each variable radius plot was cal- 
culated [III]. The ground flora species were assessed 
within a circular 4-m? plot, with each species being 
given a percentage cover value to the nearest 10%, 
with a minimum value of 5% [II]. 


The TWINSPAN classification and the recognition of 
five forest-types 

The TWINSPAN classification of 1127 of the 1200 
ground flora plots (based on the species composition 
and percentage cover of their ground flora species) 
resulted in the recognition of 11 ground flora plot 
groups or community-types [V] (see Sobey and Glen 
(2002) for a full description of the methodology — 
note that 73 plots were omitted from the analysis 
either because no trees were recorded in the area of 
the sampling point, or because no ground flora species 
were present in the 4-m* plot). These plot groups 
were then examined in terms of the species composi- 
tion of the tree canopy and other properties at the sam- 
pling sites (especially soil drainage), and as a result 
five major forest-types were recognized [VII]. (See 
Sobey and Glen (2002) for summary descriptions of 
each of these forest-types and maps showing the dis- 
tribution of their sampling points on the island.) 


2. The 1990 aerial survey: the forest stand database 
Complete aerial photographic coverage of the 
island was carried out in mid-summer 1990 as part of 


506 


1990 AERIAL 
PHOTO SURVEY: 
1:17 500-scale photos 
of the whole Island 


IV 
PHOTO-INTERPRETATION: 


demarcation and description 
of 82 957 forest stands: 
= Stand database 


XI 


THE CANADIAN FIELD-NATURALIST 


a Il 


1991 FIELD SURVEY: 
Ground flora data from 
4-m plots at 1200 
sampling points 


TWINSPAN 
CLASSIFICATION: 
=» Recognition of 
11 ground flora 
community-types 


VII ANALYSIS of 
TWINSPAN GROUPS 
with the aid of tree canopy 
and soil series data 
for the 1200 points: 
=» Recognition of five 
forest-types 


VIE SELECTION of 


TREE CANOPY & SOIL 
DRAINAGE CRITERIA 
for classification of 82 957 
stands into five forest-types 


COMPUTER SORTING 
of the 82 957 forest stands corresponding to 
the five forest-types, using the selected tree 
canopy and soil drainage criteria 


COMPUTER MAPPING 
of the stands corresponding to each 
of the island's five 1990 forest-types 


Vol. 118 


1991 FIELD SURVEY: 

Tree species data from 

‘variable-radius’ plots at 
1200 sampling points 


Vi 1988 P.E.I. 
SOIL SURVEY: 
Description and 
mapping of 44 
soil series for 
the Island 


IX DATABASE of 
SOIL SERIES & 
DRAINAGE CLASS 
for the 1200 
sampling points 


x 
DATABASE of SOIL 
DRAINAGE CLASS 
for 82 957 forest 
stands 


FIGURE 1. The stages leading to the mapping of the 1990 forest-types on Prince Edward Island: a flow diagram showing the 


databases used (in the boxes with heavy borders) and the analyses carried out. 


the 1990-1992 Prince Edward Island Forest Biomass 
Inventory [I]. The false-colour infra-red photographs 
resulting from the survey (scale c. 1:17 500) were 
analysed by trained photo-interpreters in the Forestry 
Division with the purpose of dividing the total forest- 
ed area into “stands” which were then recorded on a 
map as “polygons” [IV]. For the purpose of the 


photo-interpretation, a stand was taken to be a group 
of trees having a relatively uniform visual appearance 
from the air as evident in the aerial photograph. 
Stands could comprise a single species or several or 
many species, but they are generally of a uniform 
species composition, height and density. For each of 
the 82 957 stands, up to five tree species, each con- 


2004 


SOBEY AND GLEN: PRESENT AND PAST FOREST-TYPES OF P.E.I. 


507 


FIGURE 2. The distribution of “high forest”, based on photo-interpretation of the 1990 aerial photographic survey of the 
island. (High forest excludes Speckled Alder woods, plantations, and clear-cut and burned areas.) 


tributing greater than 5% crown closure to the canopy, 
were recorded, with the percentage contribution of 
each species being estimated to the nearest 10%. The 
database resulting from the photo-interpretation was 
computerised, and the stand boundaries were linked 
to the Geographic Information System (G.LS.) for the 
province, enabling the direct plotting onto maps of 
stand boundaries and areas, as well as the extraction 
of data on their tree species make-up. 


3. The 1988 soil survey of Prince Edward Island: a soil 
series database 

The most recent major soil survey of the island 
(MacDougall et al.1988) has resulted in the classifi- 
cation of the island’s soils into 44 soil types or “series”, 
and the mapping (scale: 1:10 000) of the distribution 
of these series over the whole island [VI]. The soil 
maps have been digitized in the G.I.S. for the island, 
such that the soil series at each of the sampling points 
in the 1991 field survey [IX], as well as for each forest 
stand polygon in the aerial stand database [X], are 
accessible by computer. For this study it was decided 
that it would also be useful to group the 44 soil series 
into six drainage classes: rapidly-drained, well- 
drained (coarse-textured parent materials), well- 
drained (medium-textured parent materials), imperfect- 
ly-drained, poorly-drained, and organic soils. 


Selection of the tree canopy and soil drainage criteria. 
The selection of the criteria for dividing the 82 957 
stands of high forest (i.e., the area shown in Figure 2) 


into the five forest-types [VII] (or “stand-types” as 
they will be termed to distinguish them from the 
TWINSPAN-defined forest-types), was based on the 
tree canopy and soil drainage properties of the five 
TWINSPAN forest-types as evident from the 1991 
field survey (Sobey and Glen 2002). Four trial com- 
puter sortings on selected criteria were carried out in 
succession; the results of each trial were evaluated 
and the criteria for each of the five stand-types were 
then redefined and a new sorting was carried out. 
Eventually the perfected criteria were chosen (Table 
1), and all of the stands in the database were comput- 
er-sorted on these criteria [XI], followed by the com- 
puter-printing of maps showing the distribution of 
each stand-type on the island [XII]. It should be 
noted that in any of the sortings the order of the sort 
(i.e., of the selecting and removal of each stand-type) 
is important, since once a stand has been sorted it is 
normally no longer available for inclusion in subse- 
quent stand-types even if it should meet their particu- 
lar criteria. The following is a more detailed discus- 
sion of the criteria used for each of the five 
forest-types, as well as of some of the changes made 
as a result of the trial sortings: 


I. Upland hardwood forest. In the TWINSPAN classi- 
fication Sugar Maple (Acer saccharum) and American 
Beech (Fagus grandifolia) had been virtually restrict- 
ed to sampling points belonging to the upland hard- 
wood forest-type (Sobey and Glen 2002). Yellow 


508 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE 1. The criteria used in the sorting of 82 957 stands of “high forest” on Prince Edward Island into five stand-types 
corresponding to the five forest-types recognized in a TWINSPAN classification of 1127 ground flora plots. 


SELECT AND REMOVE: stands designated as clear-cut, burned, plantation, windfall, or alder. This leaves 82 957 stands of 


“high forest” 


First Sort: Excluding any stands containing White Ash, Eastern White Cedar or White Elm, select all stands containing 
Sugar Maple, American Beech or Eastern Hemlock, plus all stands with Yellow Birch, but for the last species excluding 
those stands on poorly-drained, imperfectly-drained or organic soils. 


1. Upland hardwood forest 


SECOND Sort: Excluding any stands containing White Ash, Eastern White Cedar or White Elm, select all stands containing 


Black Spruce with crown closure 2 50%. 


2. Black Spruce forest 


THIRD Sort: Select all stands containing White Elm, White Ash, or Eastern White Cedar, and/or classed as ‘swampy’, plus 
all stands with a crown closure of total hardwoods = 50% and occurring on poorly-drained or imperfectly-drained soils. 


3. Wet rich woodland 


FourTH Sort: Select all stands containing White Spruce with crown closure = 40%. 


4. White Spruce woods 


RESIDUAL STANDS: 


Birch (Betula alleghaniensis) had a somewhat wider 
distribution, also occurring at a rather high level at 
wet rich woodland sampling points. In the initial trial 
sortings all stands containing any presence of Sugar 
Maple, American Beech and Yellow Birch were select- 
ed; however, for the final sorting it was decided to 
exclude those stands selected due to the presence of 
Yellow Birch that occurred on poorly-drained, imper- 
fectly-drained or organic soils, soils more characteristic 
of wet rich woodland than of upland hardwood forest. 
Also, in the final sort, Eastern Hemlock (Tsuga 
canadensis) was included as a selecting species, and 
stands containing American Elm (Ulmus americana), 
Eastern White Cedar (Thuja occidentalis) and White 
Ash (Fraxinus americana), all characteristic of wet rich 
woodland, were specifically excluded. 


2. Black Spruce forest. The problem in segregating 
stands of this forest-type was choosing a minimum 
percentage crown closure for Black Spruce. Since the 
mean percentage woody biomass contribution of 
Black Spruce at the Black Spruce forest sampling 
points was 53.1% (Sobey and Glen 2002), it was decid- 
ed to set both a high value and also one that made ecolog- 
ical sense: the criterion used in the final sort was that 
for a stand to be assigned to Black Spruce forest, at 
least 50% of its canopy had to be Black Spruce. 
From the earlier trial sortings it had also become evi- 
dent that it was necessary to exclude any stands con- 
taining American Elm, White Ash or Eastern White 
Cedar, species virtually absent from this forest-type. 


3. Wet rich woodland. In the early trial sortings an 
attempt was made to select stands of this forest-type 
on the percentage crown closure of its most impor- 
tant tree species: Red Maple (Acer rubrum) — a mini- 
mum crown closure of = 25% was set. However, the 
problem was that Red Maple was also present at high 


5. “Disturbed forest” 


levels in the still unsorted “disturbed conifer-dominated 
forest’. It was thus decided to make use of the fact 
that 80.2% of the sampling points of the TWINSPAN 
wet rich woodland occurred on imperfectly- and 
poorly-drained soil series (Sobey and Glen 2002). In 
the end the criterion used was firstly to select all stands 
containing the minor tree species diagnostic of this for- 
est-type (as evident in the TWINSPAN classification): 
ie., White Ash, American Elm and EasternWhite Cedar, 
as well as all stands classified as “swampy” by the 
photo-interpreters. Then, since an overall factor dis- 
tinguishing the TWINSPAN wet rich woodland sam- 
pling points from the disturbed conifer-dominated 
forest was a higher level of total hardwoods (Sobey 
and Glen 2002), we selected all stands with a minimum 
total hardwood crown closure of 50%, in combina- 
tion with a requirement that all such stands be on 
poorly- and imperfectly-drained soils. 

Having selected the stands that appear to be derived 
from pre-European settlement forest-types, the next step 
was to separate the remaining stands into the two types 
of successional and disturbed forest: i.e., White Spruce 
woods and disturbed conifer-dominated forest. 


4. White Spruce woods. Here it was a matter of selecting 
the minimum percentage crown closure of White 
Spruce for a stand to be assigned to White Spruce 
woods. Given the fact that the mean percentage contri- 
bution of White Spruce to the woody biomass of 
TWINSPAN White Spruce woods sampling points was 
62.2 % and in the disturbed conifer-dominated forest it 
was 16.7% (Sobey and Glen 2002), a middle level 
(40%) was chosen. 


5. Disturbed conifer-dominated forest (or simply 
“disturbed forest”). The residual stands should by 
default correspond to the remaining TWINSPAN for- 
est-type (disturbed conifer-dominated forest), even if 


2004 


SOBEY AND GLEN: PRESENT AND PAST FOREST-TYPES OF P.E.I. 


509 


TABLE 2. The number of hectares and stands (also as percentages) assigned to each stand-type in the computer-sorting. A 
percentage breakdown of the stand area by drainage class is also given, as is its status in 1935 as either forest-covered or 
cleared land as determined from a 1935 aerial photographic survey of the whole island. (In brackets after the forested per- 
centage is the equivalent parameter for the TWINSPAN forest-type sampling points of the 1991 field survey — those bolded are 


within + 20% of the stand value.) 


STAND-TYPES 


Upland Black 
Hardwood Spruce 
Forest Forest 
HECTARES 55 043 33 106 
Percentage of area Dei 13.1 
NUMBER OF STANDS 16 170 8 049 
Percentage of stands 19.5 9.7 
DRAINAGE CLASS (% of area) 
Organic 0.04 6.8 
Poorly-drained 5.8 54.8 
Imperfectly-drained 39 16.8 
Well-drained (medium) LG) 1.1 
Well-drained (coarse) 74.4 9.0 
Rapidly-drained 12.9 10.2 
Unclassified 0.28 s3 
STATUS in 1935 (% of area) 
Forested 93.8 TP 
(86.4) (67.5) 
Cleared land 6.2 22.8 


all are not in fact dominated by conifers. In the trial 
sortings an additional fifth sort was carried out in 
order to separate out the residual stands that were 
indeed conifer-dominated. This was done by sorting 
on a minimum conifer crown closure of = 50% — this 
selected 42% of the residual stands. The other group 
was thus termed “disturbed hardwood-dominated for- 
est’, i.e., comprising residual stands with hardwoods > 
50% crown closure. However, in the end, the distinc- 
tion between hardwood and softwood domination in 
the residuals was not considered important and these 
stands have been simply termed “disturbed forest”. 


Results 

Table 2 presents the results of the final computer 
sorting, expressed in terms of the area in hectares and 
the number of aerial stands assigned to each stand- 
type, both also expressed as percentages. Table 2 also 
gives a breakdown of the soil drainage classes for 
each of the stand-types and their status in 1935 (as 
“cleared land” or “forested”), as determined from 
analysis of the earliest aerial photographic survey of 
the island (see Glen 1997). The tree species make-up 
of the five stand-types is shown in Table 3 and the 
computer-printed maps showing their distribution on 
the island are shown in Figures 3 to 7. 

In the sorting, the 1990 “high forest” of the island 
was divided into five stand-types — see Appendix | for 
summary descriptions of each of these stand-types. 
These fall naturally into two groups, each of which 
occupies about half of the island’s forested area: (1) 


White 
Wet Rich Spruce Disturbed 
Woodland Woods Forest TOTALS 
42 192 57 982 65 259 253 582 
16.6 22.9 Dwi 100 
13 863 22 709 22 166 82 957 
16.7 27.4 26.7 100 
0.78 0.58 0.86 1.4 
70.1 11.6 14.5 25.8 
24.6 8.1 7.8 10.7 
0.74 2.8 5.0 2.4 
2D) 48.3 46.2 35.6 
13 213 24.1 14.9 
0.18 IED eS) 0.93 
75.8 37.6 67.4 69.0 
(79.5) (24.4) (74.2) (69.5) 
24.2 62.4 32.6 31.0 


the three stand-types which appear to be recognizable 
descendants (even if heavily-modified) of pre-European 
settlement forest-types (upland hardwood forest, 
Black Spruce forest and wet rich woodland) — these 
make up 51.4% of the area under high forest; (2) the 
two other stand-types (old field White Spruce woods 
and “disturbed forest’), representing more disturbed 
and/or successional types not likely to have been impor- 
tant in the pre-European forest — these make up the 
other 48.6%. It is likely that before the period of 
European settlement, land surfaces now under these 
two latter stand-types would have been covered by 
one of the three “primary” stand-types. 


Evaluating the validity of the stand-types 

One way of assessing the validity of the products of 
the sorting is to compare, as in Table 3, the tree species 
composition of each stand-type (in the form of per- 
centage crown closure) with the equivalent parameters 
of the corresponding TWINSPAN forest-type: the per- 
centage contribution to woody biomass in the 1200 
variable-radius plots of the ground survey, and the con- 
tribution of each of the tree species to the percentage 
crown closure in those stand polygons in which the 
1200 variable-radius plots were located. Another way 
to assess the validity of the sorting is to compare the 
drainage properties and the status in 1935 of the aerial 
stand-types with those of the TWINSPAN-based forest- 
types (Tables 2 and 4). The data required for these 
comparisons were extracted using the Database pro- 
gram “FoxPro” from the 1991 field survey database 


510 THE CANADIAN FIELD-NATURALIST Vol. 118 


TABLE 3. The mean percentage crown closure (+ S.E.) of the principal tree species in each stand-type (based on all the 
stand-polygons of each stand-type including those in which the species was not recorded). To enable comparison of these 
values with the equivalent parameters for the TWINSPAN forest-types, placed below in round brackets is the mean percentage 
contribution (+ S.E.) of each tree species to the total tree biomass in the variable-radius plots of the corresponding 
TWINSPAN forest-type; and in square brackets the mean percentage crown closure of the tree species (+ S.E.) in those stand- 
polygons in which the variable-radius plots were located. For crown closure values (unbracketed) > 10%, a bolded percent- 
age within the brackets indicates that there is “good agreement” between the two values (i.e., the bracketed value lies within 
+ 20% of the unbracketed value), while an italicized bolded percentage indicates a “notable discrepancy” between the two 
values (i.e., the bracketed value is > + 50% of the unbracketed value); where crown closures (unbracketed) were < 10%, the 
differences were assessed subjectively on the basis of their relative magnitudes. 


STAND-TYPES 


ALL STANDS 
Upland Black Wet White (All 1200 plots +) 
Hardwood Spruce Rich Spruce Disturbed [AIl 1200 sampling 
Forest Forest Woodland Woods Forest point stands] 
CONIFERS: 
Picea mariana 0.22+0.020 71.4+0.18 9.9+0.13 1.1 + 0.034 5.3 + 0.079 10.4 + 0.080 
(0.72+0.35) (53.1 +4.9) (3.8 + 1.2) (4.3 + 1.5) (10.1 + 1.4) (5.4 + 0.57) 
[0.79 + 0.28] [39.6 + 3.8] [13.3 + 1.7] [4.1 + 1.1] [12.4 + 1.3] [9.1 + 0.59] 
Picea rubens 0.59+ 0.032 0.69 + 0.042 2.9 + 0.072 0.47+0.021 7.1 +0.096 2.7 + 0.036 
(0.88 + 0.27) (17.3 +3.8) (0.51 + 0.30) (4.6 + 1.2) (Gfe Wien bi) (2.8 + 0.32) 
[0.98 + 0.28] [3:3 1:2] (5.4 + 1.1] [3.0 + 1.1] [3.6 + 0.66] [2.6 + 0.28] 
Picea glauca 4.8 + 0.086 1.6 + 0.068 8.2+0.11 68.1+0.13 10.4+0.078 24.2+0.11 
(8.6 + 0.92) (32 £723) (18.0 + 2.2) (62.2+3.4) (16.7+ 1.5) (17.0 + 1.0) 
[10.3+0.94] [25.1 + 3.4] [7.3 + 0.98] ATDE3:01 ) 7a 13) [17.5 + 0.76] 
Abies balsamea 9.0 + 0.093 4.8 + 0.086 5.6 + 0.072 3.8 + 0.053 9.0 + 0.089 6.6 + 0.037 
(18.9 + 1.2) (6.9 + 1.9) (13.5 + 1.5) (8.4 + 1.7) (24.9 + 1.6) (17.6 + 0.75) 
[10.2 + 0.60] [7-32] [7.8 + 1.0] [5.6+ 0.77] [11.1 + 0.67] [8.9 + 0.33] 
Tsuga canadensis 0.12 + 0.012 0 0 0 0 0.023 + 0.0023 
(0.32 + 0.16) (0) (0) (0.81 40.54) (0.42+0.25) (0.40 + 0.14) 
[0] [0] [0] [0] [0] [0] 
Larix laricina 0.082 +0.0097 8520.11 5.4 + 0.075 6.0 + 0.075 12.8 + 0.16 7.1 + 0.052 
(0.48 + 0.18) (9.0+2.1) (7.3 + 1.7) (3.7+0.84) (3.84 0.69) (2.6 + 0.31) 
[1.5 + 0.35] [9.5 + 1.5] [7.8 + 1.2] [8.8 + 1.6] [7.6 + 0.81] [5.3 + 0.37] 
Thuja occidentalis 0 0) 1.2 + 0.037 0 0 0.20 + 0.0064 
(0.42+0.25) (0.12 + 0.13) (TET) (0) (0.64+0.32) (0.93 + 0.23) 
[0.023 + — ] [0] [0.38 +— ] [0.15+-] [0.39+0.14] [0.17 + 0.041] 
TOTAL CONIFERS 14.9 87.3 34.9 80.4 46.3 51.4 
(30.4) (94.4) (50.0) (84.1) (64.5) (47.3) 
[23.8] [84.7] [42.0] [69.6] [52.4] [44.4] 
BROAD-LEAVES: 
Acer rubrum Sysifas OBL 4.8 + 0.086 25) 210a3 5.4 + 0.061 20.1 + 0.12 18.7 + 0.063 
(33.3+41.4) (2.5 +0.77) (24.6 + 2.4) (4.6 + 1.1) (19.0 + 1.5) (Bales 1) 
[30.4 + 0.73] [5.1 + 1.1] [26.3 + 1.5] [8341.1] [18.9+0.92] [21.7+0.52] 
Acer saccharum 21.6+0.10 0 0.45 + 0.030 0 0 4.3 + 0.036 
(9.7 + 0.89) (0) (1.7+0.56) (0.59+0.46) (0.28 +0.13) (4.9 + 0.54) 
[16.7 + 0.76] [0.5 +-] [2.1 + 0.55] [20160] [35.053] [7.7 + 0.38] 
Betula alleghaniensis 7.8 + 0.070 0) 0.22+0.016 0.013 + 0.0026 0.090 + 0.0024 1.6+0.017 
(5.6 + 0.62) (0) (3.3 + 0.84) (0.22+0.13) (1.4+0.40) (3.9 + 0.39) 
[6.7 + 0.41] [0] (0.71+0.22] [1.0+0.35] [0.92 + 0.23] (3.0 + 0.19] 
Betula papyrifera 137+ 01087, 2:5 = 01059 12.7 + 0.089 6.3+0.059 13.4+0.088 10.3 + 0.039 
(9.2+0.74) (0.63 + 0.36) (5.6 + 0.94) (3.6+0.78) (41+0.60) (7.3 + 0.50) 
{13.1+ 0.53] [2.5 + 0.65] (11.9+0.97] [6.9+0.85] [11.4+0.66] [10.8 + 0.33] 
Fagus grandifolia 1.5 + 0.036 0 0.017 + 0.0044 0 0 0.30 + 0.0073 
(4.1 + 0.58) (0) (0) (0.37+0.24) (0.23+0.11)  (2.7+0.37) 
[1.1 + 0.19] [0] [0.19 +—] (0.074+-—] [0.21+0.10] [0.49 + 0.074] 


(continued) on next page 


2004 SOBEY AND GLEN: PRESENT AND PAST FOREST-TYPES OF P.E.I. S51] 


TABLE 3. (continued from previous page) 


STAND-TYPES 


ALL STANDS 


Upland Black Wet White (All 1200 plots +) 
Hardwood Spruce Rich Spruce “Disturbed [AIl 1200 sampling 
Forest Forest Woodland Woods Forest” point stands] 
Populus spp. ¢ 4.6 + 0.075 5.0 + 0.090 21.4+0.15 7.6+0.070 18.8+0.14 I23:2.0/057 
(4.1 + 0.65) (1.1 + 1.0) (7.8 + 1.6) (4.7 + 1.2) (7.4 + 1.1) (6.6 + 0.62) 
[6.9 + 0.53] [6.8 + 1.2] {11.2+1.0] [10.7+1.1] [10.9+0.80] [9.6 + 0.39] 
Fraxinus americana 0 0 0.32 + 0.019 0 0 0.053 + 0.0031 
(0.40 + 0.21) (0) (1.6 + 0.75) (0) (O) (0.46 + 0.16 ) 
[0] [0] [0] [0] [0] [0] 
Ulmus americana 0 0 0.84 + 0.035 0 0 0.14 + 0.0058 
(0.26 + 0.14) (O) G2 saie1) (0) (0) (0.57 + 0.17) 
[0.47+ — | [0] (0.13 +-] [0] [0.035+-—] [0.042 + 0.019] 
Alnus incana 0.047 + 0.011 0.26 + 0.027 4.8+0.11 201039" Os 01052 6.4 + 0.071 
(-) (-) (-) (-) (-) (-) 
[0.16 + 0.12] [0] (4.0+1.2] [1.1+0.63] [0.85 + 0.38] [et e10:22] 
TOTAL BROAD-LEAVES 85.1 12.5 65.1 19.6 3 47.8 
(excluding Alnus) (70.0) (5.3) (50.0) (16.5) (35.0) (52.7) 
[75.6] [14.9] [56.7] [30.3] [46.9] [55.6] 
Number of Stands 16,170 8,049 13,863 22,709 22,166 82,957 


¥ For “all 1200 plots” the value in round brackets is the percentage contribution to the total above ground biomass (oven- 
dry tonnes per hectare) in all 1200 variable-radius plots. 

+ Because it is impossible to distinguish the three native poplar species in aerial photographs (Populus tremuloides, P. 
balsamifera, P. grandidentata) they were grouped as “Populus spp.” — however almost all of this is likely to have been 
P. tremuloides (Trembling Aspen). 


FIGURE 3. The distribution of stands of upland hardwood forest in 1990. 


a2 THE CANADIAN FIELD-NATURALIST Vol. 118 


Ficure 4. The distribution of stands of Black Spruce forest in 1990. 


FicureE 5. The distribution of stands of wet rich woodland in 1990. 


2004 


SOBEY AND GLEN: PRESENT AND PAST FOREST-TYPES OF P.E.I. 


FIGuRE 6. The distribution of stands of White Spruce woods in 1990. 


FIGURE 7. The distribution of stands of “‘disturbed forest” in 1990. 


514 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE 4. The soil drainage classes of the stand-types (derived from the 1:10 000 soil series maps of MacDougall et al. 
(1988)) expressed as the percentage of the stand-type’s total area in each drainage class, ignoring the area of unclassified 
drainage. These are compared with the drainage classes of the corresponding TWINsPAN forest-types: (1) (in round brackets) 
the soil drainage class at the sampling points of the 1991 field survey as determined from soil pits, and (2) [in square brack- 
ets] the drainage class of those soil series polygons in which the sampling points were located, derived also from Mac- 
Dougall et al. (1988), both values being expressed as the percentage of the total area or of the sampling points in each 


drainage class. * 


STAND-TYPES 


Upland Black 
Hardwood Spruce 
SOIL DRAINAGE CLASS Forest Forest 
Poorly- and imperfectly- 9.8 79.4 
drained soils (+ organic)(%) (1.9) (66.3) 
[12.9] [73.8] 
Well- and rapidly- 90.2 20.6 
drained soils (%) (98.1) (33.8) 
[87.1] [26.3] 


White ALL 
Wet Rich Spruce “Disturbed STANDS 
Woodland Woods Forest” —_ (or points) 
95.7 20.6 23.6 39.2 
(58.3) (1.5) (15.9) (18.4) 
[80.3] [14.0] [37.5] [33.6] 
4.3 79.4 76.4 60.8 
(41.6) (98.5) (84.1) (81.6) 
[19.9] [85.9] [62.5] [66.4] 


*For the meaning of the bolding and italicizing see the caption of Table 3. 


and the 1990 aerial stand database, and then linked to 
the “MapInfo” G.LS. for the province. 


(1) Comparing the composition of the tree canopy 

To facilitate the overall interpretation of the many 
comparisons being made, indices were selected for 
assessing the closeness of the two values: if the equiv- 
alent parameter for the TWINSPAN-based forest-type 
lay within + 20% of the aerial-stand value, this was 
taken to indicate “good agreement” between the two 
values, whereas if the parameter was greater than + 50% 
of the aerial-stand value this was considered a “notable 
discrepancy”. On the basis of these criteria, it is evident 
that in overall terms there is a reasonably good fit 
between the mean tree canopy composition of the 
stand-types based on all of the stand polygons (Table 
3), and the canopy composition of those stand poly- 
gons that contained the 1200 variable-radius plots: there 
are 29 “good agreements” and only two “notable dis- 
crepancies” for the crown closure of individual species. 
However, when the mean tree canopy composition for 
all of the stand polygons is compared with the per- 
centage contribution to woody biomass in the variable- 
radius plots, the match is not as good: here there were 
15 “good agreements” and 17 “notable discrepancies”. 
This greater level of discrepancy is not surprising 
considering that this latter comparison is between 
plots of different size (i.e., forest stands of variable size, 
but measured mostly in hectares (the mean stand area 
was about 3 hectares) and variable-radius plots with a 
mean size of about 100 m7), as well as different parame- 
ters (percentage crown closure as determined from an 
aerial photograph, with the percentage contribution to 
woody biomass based on the trunk diameter of the 
trees within the variable-radius plots). 

Also relevant here is the greater difficulty in identi- 
fying tree species from an aerial photograph compared 


with the examination of actual specimens in a field 
survey. The photo-identification was based on such 
characters as texture, shape, colour and site (e.g., wheth- 
er on upland or lowland). The conifers in particular 
presented problems: Balsam Fir (Abies balsamea) when 
present as an understorey species may not be apparent 
from the air; the three spruce species were frequently 
identified from their site: i.e., if in upland forest, it 
was listed as Red Spruce (Picea rubens), in lowland, 
Black Spruce, and on old-field sites, White Spruce, but 
this was not an infallible approach. Thus the photo- 
identification was not 100% accurate: the standard re- 
quired for the 1990 aerial survey, confirmed by ground 
checks, was that at least 70% of the species-cover 
identification in a stand be correct for at least 90% of 
the time. Bearing these points in mind we observe 
the following: 


Upland hardwood stands. In almost all of the compar- 
isons (Table 3) there is generally a good correspondence 
between the parameters of the aerial stand-type and 
those of the TWINSPAN-defined forest-type. 


Black Spruce stands. There are problems with the 
spruces (perhaps due to the problem in identifying the 
spruces, discussed above): in Table 3 the most notable 
discrepancy is the very low Red Spruce contribution 
to the aerial stand-type (<1%) compared with its level 
in the variable-radius plots (17%). Also evident is the 
fact that Black Spruce is at a somewhat higher level in 
the aerial stand-type (71%) compared with either of the 
values for the TWINSPAN-defined forest-type. This is 
presumably due to the high crown closure level (i.e., 
50%) that was set as the criterion for inclusion of Black 
Spruce stands in this group. 


Wet rich woodland stands. The chief species, Red 
Maple, shows no difference in either of the comparisons 


2004 


(Tables 3). However, there are some discrepancies: the 
use of a sorting criterion of 2 50% hardwoods must be 
the reason for the higher overall hardwood contribution 
in the aerial stand-type compared with the TWINSPAN 
forest-type, as well as the higher individual contribu- 
tions of Trembling Aspen (Populus tremuloides) and 
White Birch (Betula papyrifera), though there is no dis- 
crepancy for the latter species in the comparison of the 
crown closures. Concomitantly, the conifer contribution 
is less in the aerial stand-type than in the variable-radius 
plots, but again there is less discrepancy for the crown 
closure comparison. 


White Spruce stands. In overall terms there is a good 
correspondence between the two classifications, with 
the main species, White Spruce, giving a reasonably 
good fit — this time the better fit is for the variable- 
radius plot comparison (Table 3). 


“Disturbed forest” stands. This residual group has a 
smaller total conifer component than the TWINSPAN 
“disturbed conifer-dominated forest” (this discrepancy 
is less evident in the comparison of the crown closures 
— Table 3), and this is reflected especially in the levels 
of Balsam Fir (for the variable-radius plot comparison 
only), and to a smaller degree, in White Spruce and 
Black Spruce. Oddly, Tamarack (Larix laricina) is con- 
siderably higher in the aerial stand-type than it is in the 
variable-radius plots. Concomitantly, the hardwoods 
(notably Trembling Aspen) are higher in the aerial 
stand-type, while there is no difference for the levels of 
Red Maple. 


(2) Comparing the soil drainage classes and the 1935 
Status 

There is a reasonably good agreement between the 
aerial stand-types and the TWINSPAN forest-types 
in their drainage class breakdown (Table 4), especially 
for the drainage classes as determined from the soil 
series at the 1200 sampling points. There is less agree- 
ment with the drainage classes as determined from the 
soil pits dug at the sampling points during the field 
inventory. The one showing the least correspondence is 
the wet rich woodland which has higher levels of poor- 
ly- and imperfectly-drained soils in the aerial stands, 
due to the direct use of these drainage classes as a 
sorting criterion. The status of the stands and sam- 
pling points in 1935 (1.e., whether they were forested 
or cleared land at that time) is in broad agreement for 
most of the forest-types (Table 2), with the greatest 
discrepancy being for the White Spruce woods. 


Discussion 

The TWINSPAN analysis of the 1991 field survey 
data (Sobey and Glen 2002) indicated the presence of 
five main forest-types on Prince Edward Island, three 
of these showing a relationship with pre-European 
settlement forest-types and two appearing to largely 
owe their presence to human influences. The objective 
of the current paper was the production of maps show- 


SOBEY AND GLEN: PRESENT AND PAST FOREST-TYPES OF P.E.I. 


S15 


ing the total distribution of these five forest-types. This 
objective has been achieved, and for the first time ever 
we have fine-scale whole-island maps showing the 
distribution of the main forest-types. Furthermore, these 
maps have been produced using largely objective cri- 
teria and methods, and they are based on the actual for- 
est present at a specific time (in the summer of 1990), 
with every mapped stand being locatable on the ground 
and thus potentially verifiable. 

How valid are the maps as a picture of the distribu- 
tion of the five forest-types on the island? Firstly, the 
maps appear to make ecological and geographical sense: 
the upland hardwood forest (Figure 3) occurs in areas of 
higher elevation (in island terms), and on soils having 
good drainage (Tables 2 and 4). In such areas the map- 
ping indicates that it is present either as small woodlots 
surrounded by farmland, or as larger connected parcels 
(at the back end of farms) — especially in the central 
and south-eastern hill-lands, where steeper slopes and 
shallow soils placed restrictions on forest clearance 
in the past. 

It is also evident that the Black Spruce forest and the 
wet rich woodland, which occur primarily on soils with 
poorer drainage (Tables 2 and 4), are segregated geo- 
graphically from the upland hardwood forest. These 
predominate in areas of lower elevation, notably in parts 
of the east and west of the island (Figures 4 and 5). 
Though both of these forest-types occur in the same 
geographical areas, the TWINSPAN analysis indicated 
that they have very different ground and tree vegetation 
(Sobey and Glen 2002). They also have very different 
soil chemical properties — the wet rich woodland has 
soils that are comparatively base-rich and of higher 
pH with a lower C/N (carbon-nitrogen) ratio, than the 
soils of Black Spruce forest, more acid and base-poor 
with a higher C/N ratio. This dissimilarity was also 
evident from the fact that there was no overlap in 
their graphical distributions in the DECORANA ordi- 
nation (Sobey and Glen 2002). Given their similar geo- 
graphical distribution on the island, the question is 
whether in the mapping we have been able, using tree 
and soil criteria alone, to separate the stands of these 
two wet forest-types from each other. The differences in 
their tree composition parameters, evident in Table 3, 
suggest that we have. 

Stands of the other two forest-types, the White Spruce 
woods and the disturbed forest, have a wider and more 
scattered distribution (Figures 6 and 7), occurring 
less as larger connected parcels, than as individual small 
stands, which is consistent with the fact that their occur- 
rence is largely the product of human activity, which 
will tend to occur at a scale determined by the own- 
ership of blocks of land. At the same time there is a 
geographical aspect to their distribution, with the White 
Spruce woods prevalent in the central hill-lands and 
in the eastern part of the island, its spatial distribution 
evidently determined by the geography of field and 
farm abandonment, while the disturbed forest is con- 


516 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


TABLE 5. For each soil drainage class, the total area (in hectares) under the three ‘primary’ forest stand-types in 1990, and 
the percentage of this area assigned in the computer-sorting to each stand-type is shown. 


DRAINAGE CLASS 


PRIMARY FOREST Poorly- _ Imperfectly Well-drained Well-drained 

STAND-TYPES Organic drained drained (medium textured) (coarse textured) Rapidly-drained 
Upland Hardwood Forest 0.88 6.2 12.0 68.7 91.2 64.4 
Black Spruce Forest 86.5 35.6 30.7 16.7 6.7 30.6 

Wet Rich Woodland 12.6 58.2 57:3 14.6 2a 5.0 

Total Hectares 2 620 50 912 18 089 Zl Si 44 865 11 073 


centrated largely in the far west and in the east of the 
island. 

Although a significant and meaningful ecological 
picture has emerged from the sorting and mapping, at 
the same time it is important to point out some limi- 
tations: a basic one is that any sorting of forest stands 
into forest-types is a classification procedure, and any 
process of classification is a simplification. Compound- 
ing this was the fact that the stand sorting was reliant 
on the products of three other classification procedures: 
TWINSPAN, stand delineation (strictly-speaking not 
a classification procedure, but involving demarcation 
of the forest area into individual stands), and the soil 
series classification for the province. Each of these 
three prior procedures will have their own particular 
limitations. 

It should also be pointed out that although three cen- 
turies of forest destruction and exploitation by Euro- 
peans have undoubtedly generalized and simplified 
the island’s forest cover, the five maps are still a sim- 
plification of what was present in 1990. As noted in 
Sobey and Glen (2002) there are likely to have been 
other forest-types present but so uncommon as not to 
have been picked up in the 1991 field survey in suffi- 
cient quantity to be recognized as separate types (e.g., 
areas under Hemlock, White Pine, Eastern White Cedar 
and White Ash, now occurring as only a few small 
stands, but likely to have been more widespread in the 
past). Also, each of the five forest-types that has been 
recognized may have had sub-types that have been 
overlooked: e.g., the TWINSPAN classification sub- 
divided the Black Spruce forest plots into wetter and 
dryer variants (Sobey 1995), but an attempt in the sort- 
ing to separate stands of these on their tree canopy com- 
position was not successful. The upland hardwood 
may also have variants dominated by particular tree 
species. The only solution to this problem of both rare 
and more refined forest-types, is firstly to define them, 
and then to locate them on the ground and map them 
separately. 

It should also be noted that the maps produced 
present a picture of the forest as it was at one particular 
time: in the summer of 1990 at the time of the aerial 
photographic survey. But the forests of Prince Edward 
Island, like forests the world over, are not fixed and 


static. Apart from natural factors, they are subject to the 
effects of continual human interference, for example, 
clear- and partial-cutting, forest clearance for agricul- 
ture, fire, and farm abandonment, as well as the natural 
successional processes resulting from these effects. 
On the other hand, although the three pre-European 
settlement woodland types may vary over time in the 
amount and proportion of the island’s area that they 
occupy, we should not expect the general pattern of 
their distributions to change, related as this is to innate 
soil properties. However, the two successional forest- 
types (White Spruce woods and disturbed forest) are 
likely to be only transitional on a site, and we may 
presume that given sufficient time, most of their sites 
would develop into one of the primary woodland types. 


Mapping the pre-European settlement forest of the 
island 

Despite the above limitations in the methodology, 
it is of interest to generalize the results even further by 
considering the potential for extending the mapping 
to areas without forest cover in 1990, incidentally, an 
aspect of all earlier attempts at mapping the province’s 
forests (Stilgenbauer 1929; Halliday 1937; Rowe 1959; 
Loucks 1962). Such a mapping may have either of 
two objectives: (1) the reconstruction of the type of 
forest likely to have occurred before forest clearance, 
i.e., the “original-natural” forest sensu Peterken (1996); 
and/or (2) the prediction of the type of forest that would 
ultimately occur over the whole island if all areas were 
allowed to revert naturally to climax forest in the future; 
i.e., the “future-natural” forest sensu Peterken (1996). 
Either of these objectives involves trying to predict the 
climax forest-type on the land area now occupied by 
White Spruce woodland and the “disturbed forest’, as 
well as the forest cover on land which in 1990 was clear 
of forest, i.e., the non-forested area evident in Figure 2, 
comprising some 43% of the island’s land area 
(Anonymous 1992). 

The easiest approach to such a mapping is to take 
advantage of the close relationship existing between 
natural vegetation and soils, by utilizing data derived 
from the comprehensive and fine-scale maps produced 
by the Prince Edward Island Soil Survey (MacDougall 
et al. 1988). We need first to examine any associations 


2004 SOBEY AND GLEN: PRESENT AND PAST FOREST-TYPES OF P.E.I. S197, 


FIGURE 8. The conjectured distribution of upland hardwood forest prior to European settlement — based on combining the 
distribution of upland hardwood stands in 1990 with that of well-drained soils as mapped by the Soil Survey of 
Prince Edward Island (MacDougall et al. 1988). 


FIGURE 9. The conjectured distribution of wet rich woodland and Black Spruce forest prior to European settlement — based 
on combining the distribution of stands of these two forest-types in 1990 with the distribution of imperfectly- and 
poorly-drained soils (excluding stream complexes) as mapped by the Soil Survey of Prince Edward Island (Mac- 
Dougall et al. 1988). 


518 


occurring between the three pre-settlkement forest- 
types and the soils of different drainage properties: 
Table 5 shows for each drainage class the percentage 
of its “primary forest’ component contributed by each 
of the three primary forest-types. 

From Table 5 it is evident that well-drained coarse- 
textured soils (and to a less extent well-drained medium- 
textured soils) have a strong association with upland 
hardwood forest (some 91% of the well-drained coarse 
soils and 69% of the medium soils have such a forest 
cover). By contrast, poorly and imperfectly drained 
soils are strongly associated with the two other pre- 
settlement forest types (wet rich woodland and Black 
Spruce forest) — some 94% of poorly-drained and 
88% of imperfectly-drained sites carry such forests. 
However, although these two “wet” forest-types (as 
noted in Sobey and Glen (2002) and above), occupy 
soils of markedly differing chemical properties (in 
terms of pH, base status and C/N ratios), such chemi- 
cal properties were not criteria used to characterize 
the soil series in the 1988 Soil Survey, and thus the 
soil series maps cannot be used to differentiate these 
two forest-types. Then there is the problem presented 
by the remaining soil drainage class, rapidly-drained 
soils (Table 5): 64% of such soils carry upland hard- 
wood forest, but a significant proportion (30%) has a 
cover of Black Spruce forest, more likely the dryer vari- 
ant of the Black Spruce forest, shown in the TWINSPAN 
analysis (Sobey 1995) to have some association with 
rapidly-drained soils. 

On the basis of the above relationships, Figure 8 is 
an attempt to show the approximate distribution of 
the pre-settlement upland hardwood forest, and pre- 
sumably also the area that is capable of reverting to 
such forest in future. The map has been constructed by 
taking the 1990 distribution of upland hardwood forest 
(Figure 3) and adding to it the area occupied by well- 
drained soils (both coarse and medium-textured parent 
materials). It is likely that additional areas under rapid- 
ly-drained soils, particularly in the south-eastern hill- 
lands, would also have carried some form of upland 
hardwood forest, but given the present state of our 
information, it is not possible at this stage to distinguish 
the hardwood-bearing areas of such soils from the 
Black Spruce areas. 

Figure 9 is an attempt to show the area occupied 
before European settlement by the two wet forest-types 
(the wet rich woodland and the Black Spruce forest). 
It has been constructed by adding to the 1990 distri- 
bution of these forest-types (i.e., Figures 4 and 5), the 
combined distribution of poorly- and imperfectly- 
drained soils. Given the present state of the informa- 
tion incorporated within the G.I.S. database for the 
province, this is about as far as we can go for wet wood- 
land. Distinguishing between these two woodland types 
would require data on the chemical properties of the 
soils. And, as noted above, Table 5 indicates that Black 
Spruce forest (presumably the dryer variant) will have 
also occurred on land having rapidly-drained soils. It is 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


also likely that it is on such soils that the now largely 
extirpated pine forests of the island would have occurred 
in the past. 

The picture that emerges from these maps is of an 
island that was mostly covered by upland hardwood 
forest, which formed a large continuous block in the 
central part of the island, with smaller but significant 
blocks in the western and eastern parts. By contrast, 
the wetter forest-types occurred at lower elevations in 
specific areas in the west and in a more scattered mosaic 
pattern in the east. It is thus on these lowland sites 
that the boreal element of the island’s forests, in the 
form of Black Spruce forest, had a greater occurrence. 
Such a division of the landscape, based on soil mois- 
ture levels, between the northern hardwood forest and 
the boreal forest, appears to be typical in the boreal- 
broadleaf ecotone (Pastor and Mladenoff 1992; Scott 
1995). 

It should be noted that compared with earlier attempts 
at mapping the pre-settlement forest (e.g., Stilgenbauer 
1929; Loucks 1962) the maps presented here have a 
much finer scale of detail in their boundary lines. 
Also, the validity of the maps, as showing the forest 
distribution before European settlement, can be test- 
ed by comparing them with early historical descrip- 
tions of the forest, especially from the data contained 
in early maps and surveyors’ field notebooks in the 
provincial archives. Not only would such a compari- 
son allow us to ascertain the forest-types that actually 
occurred on many sites before forest clearance, it 
may also provide data on the fine-scale differences in 
the forests of such sites, in some cases even as to par- 
ticular tree species. However this is beyond the scope 
of the present study. By contrast, the validity of the 
maps as pictures of the “future-natural” forest will only 
be testable from what happens if such areas revert to 
forest in future. 


Acknowledgments 

Doug Sobey thanks Professor Gerry McKenna, then 
Dean of Science at the University of Ulster, for en- 
abling him to take six months leave from University 
duties in 1996 so that he could continue his research 
into the forests of Prince Edward Island, and the P.E.I. 
Department of Agriculture and Forestry, Forestry Divi- 
sion, for allowing full access to the 1990-1992 Forest 
Inventory data. He also thanks Harry Baglole of the 
Institute of Island Studies at the University of Prince 
Edward Island for supporting the continuance of the 
research associateship which enabled the use of library 
facilities at the University. Both authors thank Delmar 
Holmstrom of Agriculture Canada, Charlottetown, for 
providing advice on the grouping of the soil series 
into drainage classes. 


Literature Cited 

Anonymous. 1992. 1990/1992 Prince Edward Island forest 
inventory: Summary. Forestry Branch, P.E.I. Department 
of Energy and Forestry, Charlottetown, Prince Edward 
Island. 


2004 


Erskine, D. S. 1960. The plants of Prince Edward Island. 
Publication 1088. Canada Department of Agriculture. 
Glen, W. M. 1997. Prince Edward Island 1935/1936 forest 
cover type mapping. Forestry Division, P.E.I. Department 
of Agriculture and Forestry, Charlottetown, Prince Edward 

Island. 

Halliday, W. E. D. 1937. A forest classification for Canada. 
Canada Department of Mines & Resources, Ottawa. 

Loucks, O. L. 1962. A forest classification for the Maritime 
Provinces. Proceedings of the Nova Scotia Institute of 
Science 25: 85-167. 

MacDougall, J. L., C. Veer, and F. Wilson. 1988. Soils of 
Prince Edward Island. Agriculture Canada, Charlotte- 
town, Prince Edward Island. 

Pastor, J., and D. J. Mladenoff. 1992. The southern boreal- 
northern hardwood border. Pages 216-240 in A systems 
analysis of the global boreal forest. Edited by H. H. Shugart, 
R. Leemans, and G. B. Bonan. Cambridge University Press, 
Cambridge, England. 

Peterken, G. F. 1996. Natural woodland — ecology and con- 
servation in northern temperate regions. Cambridge Uni- 
versity Press, Cambridge, England. 


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519 


Rowe, J. S. 1959. Forest regions of Canada. Canada Depart- 
ment of Northern Affairs and Natural Resources, Forestry 
Branch, Bulletin 123 [Reprinted 1972 with minor revisions 
as: Publication No. 1300, Canadian Forestry Service, Depart- 
ment of the Environment, Ottawa]. 

Scott, G. A. J. 1995. Canada’s vegetation — a world perspec- 
tive. McGill—Queen’s University, Montreal and Kingston. 

Sobey, D. G. 1995. Analysis of the ground flora and other 
data collected during the 1991 Prince Edward Island forest 
inventory. II. Plant community analysis. Prince Edward 
Island Department of Agriculture, Fisheries and Forestry, 
Charlottetown, Prince Edward Island. 

Sobey, D. G., and W. M. Glen. 2002. The forests of Prince 
Edward Island: A classification and ordination using multi- 
variate methods. Canadian Field-Naturalist 116: 585-602. 

Stilgenbauer, F. A. 1929. The geography of Prince Edward 
Island. Ph.D. dissertation, University of Michigan, Ann 
Arbor, Michigan. 

Watts, S. B. 1983. Forestry handbook for British Columbia 
(Fourth edition). University of British Columbia, Vancou- 
ver, British Columbia. 


Received 20 May 2003 
Accepted 4 November 2004 


APPENDIX 1. Summary descriptions of the five forest stand-types arising from the computer- 


sorting of 82 957 forest stands. 
UPLAND HARDWOOD FOREST 16 170 stands (19.5%); 

55 043 ha (21.7%) 

A stand-type of widespread occurrence on well-drained 
soils (77% of its area is on such soils — plus an additional 
13% on rapidly-drained soils), especially away from the 
coast and further back from roads, in the central and south- 
eastern hill-lands, in the north-east peninsula, and in small- 
er specific areas in the east and west of the island. 94% of 
its area had a forest cover in 1935. The tree canopy is domi- 
nated by hardwoods (accounting for 85% of crown closure) 
— particularly Red Maple (36%), with a notable contribution 
by Sugar Maple (22%), and smaller amounts of Yellow Birch 
(8%) and American Beech (2%). There is also a significant 
component of White Birch (14%) and Trembling Aspen (5%). 
The minority conifer component (15%) consists largely of 
Balsam Fir (9%), with some White Spruce (5%). The high 
contribution to crown closure by Red Maple, and the pres- 
ence of White Birch and Trembling Aspen, are indicative of 
a high level of past, and we may presume, continuing, human 
disturbance. This stand-type is the direct descendant of the 
pre-European settlement upland hardwood forest postulated 
by Erskine (1960), though subject to varying levels of past 
and current human disturbance. 
BLACK SPRUCE FOREST 8 049 stands (9.7%): 

33 106 ha (13.1%) 

A rather localized stand-type occurring primarily on areas 
with poor and imperfect drainage — 78% of its area is on 
such soils — especially in specific parts of the west and east 
of the island. The tree canopy is dominated by conifers (com- 
prising 87% of crown closure), above all Black Spruce (71%) 
— with small contributions by Tamarack (9%) and Balsam 
Fir (5%). The 13% contributed by hardwoods comes largely 
from Red Maple (5%) and Trembling Aspen (5%). Most of 


the area (77%) had a forest cover in 1935. This ‘stand-type 
corresponds to the pre-settlement lowland Black Spruce 
forest of Erskine (1960). 
WET RICH WOODLAND 13 863 stands (16.7%); 
42 192 ha (16.6%) 

A stand-type almost completely confined to areas of poorly- 
and imperfectly-drained soils (95% by ground area) espe- 
cially in the west of the island, as well as parts of the east. 
The tree canopy is dominated by hardwoods (making up 
65% of crown closure), but there is also a notable conifer 
contribution (35%). The chief hardwood species are Red 
Maple (26%), Trembling Aspen (21%) and White Birch 
(13%). (White Cedar, American Elm and White Ash are 
present at low levels). The main conifers are Black Spruce 
(10%), White Spruce (8%), Balsam Fir (6%) and Tamarack 
(5%). 76% of its area was under forest cover in 1935. This 
stand-type appears to be a derivative of the pre-settlement 
lowland Red Maple forest of Erskine (1960) — though with 
considerable effects due to cutting and other disturbances. 


22 709 stands (27.4%): 
57 982 ha (22.9%) 

A widely scattered stand-type occurring as small often field- 
shaped parcels on well- and rapidly- drained soils (78% by 
area is on such soils). It is prevalent in the central hill-lands 
and even more towards the east of the island. A high pro- 
portion of its area (62%) was cleared land in 1935. The tree 
canopy is dominated by a single species, White Spruce, at a 
high level of crown closure (68%). The remaining species 
contribute only small amounts: Trembling Aspen (8%), 
White Birch (6%), Tamarack (6%), Red Maple (5%) and 
Balsam Fir (4%). This stand-type corresponds to the succes- 
sional “old field” White Spruce woods of Erskine (1960). 


WHITE SPRUCE Woops 


520 


“Disturbed Forest’ 22 166 stands (26.7%): 
65 259 ha (25.7%) 

This residual stand-type (i.e., that left over after the stands 
of the other four types had been selected) is widely scattered, 
generally as small individual parcels (rather than large blocks), 
especially in the east and west of the island, notably on well- 
and rapidly-drained soils (75% by area occurred on such 
soils). The tree canopy has a slight predominance of hard- 
woods (54%) — the principal species being Red Maple (20%), 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Trembling Aspen (19%) and White Birch (13%). Its large 
conifer component (46%) is made up of Tamarack (13%), 
White Spruce (10%), Balsam Fir (9%), Red Spruce (7%) and 
Black Spruce (5%). 33% of its area was cleared land in 1935. 
This stand-type does not equate with any of Erskine’s pre- 
European forest-types — its species make-up and distribution 
suggest rather that it is largely a successional community 
heavily affected by past and continuing human exploitation. 


Inventaire printanier d’une frayere multispécifique : l’ichtyofaune des 
rapides de la riviere Gatineau, Québec 


ANNIE Comto!s!, FRANCOIS CHAPLEAU!, CLAUDE B. RENAUD?, HENRI FOURNIER*, BRENT CAMPBELL’, 
et RICHARD PARISEAU? 


'Département de biologie, Université d’ Ottawa, Ottawa, Ontario KIN 6N5 Canada 

Division de la recherche, Musée canadien de la nature, Casier Postal 3443, Succursale D, Ottawa, Ontario K1P 6P4 Canada 

3Direction de l’aménagement de la faune de I’ Outaouais, Société de la faune et des parcs du Québec, Gatineau, Québec J8Y 3R7 
Canada 


Comtois, Annie, Frangois Chapleau, Claude B. Renaud, Henri Fournier, Brent Campbell, et Richard Pariseau. 2004. Inventaire 
printanier d’une frayére multispécifique : ichtyofaune des rapides de la riviere Gatineau, Québec. Canadian Field- 
Naturalist 118(4): 521-529. 


Un inventaire ichtyologique printanier a été réalisé dans la riviere Gatineau, dans le premier rapide a l’amont de son embou- 
chure dans la riviere des Outaouais, pour établir la chronoséquence d’utilisation du site par les especes s’y reproduisant, dont trois 
espéces de chevaliers. L’échantillonnage des rapides a permis de capturer 2388 poissons répartis en 13 familles et 39 espeéces, 
dont six ayant fait ’ objet d’une évaluation par le COSEPAC. Seize espéces étaient en état de se reproduire lors de |’ échantil- 
lonnage. Nous avons observé une reproduction plus ou moins simultanée pour le Chevalier blanc et le Chevalier rouge, suivie 
de celle du Chevalier de riviére. Egalement, pour ces trois espéces, nous avons observé la capture des miles de stade V plus 
tot que les femelles, et nous les retrouvons en plus grand nombre. Ceci pourrait s’expliquer du fait que les males précedent 
les femelles sur les frayeéres, que celles-ci sont reconnues pour rester tres peu de temps sur les sites de fraie et qu’un 
minimum de deux males par femelle est requis pour la reproduction. Cette étude démontre que les rapides de la riviére 
Gatineau doivent étre protégés puisqu’ils sont habités par trois especes qui ont obtenu un statut spécial du COSEPAC 
(Menacée: Fouille-roche gris; situation préoccupante : Lamproie du nord et Chevalier de riviere) en plus de constituer une 
frayére multispécifique importante. 


Mots-clés: Rapides, Riviere Gatineau, Outaouais, Fraie, Chevalier blanc, Moxostoma anisurum, Chevalier de riviére, Moxostoma 
carinatum, Chevalier rouge, Moxostoma macrolepidotum, Lamproie du nord, [chthyomyzon fossor, Fouille-roche gris, 
Percina copelandi. 


A spring survey of fishes was conducted in the Gatineau River (Quebec), in the first rapids upstream of its confluence with 
the Ottawa River, to establish the fish spawning chronosequence, especially for the three species of redhorses. The sampling 
resulted in the capture of 2388 fish belonging to 13 families and 39 species. Six of these species have been assessed by 
COSEWIC. Sixteen species were in spawning readiness. Spawning of the Silver Redhorse and Shorthead Redhorse occurred 
first, followed by the River Redhorse. Stage V males of the three redhorse species were captured earlier and in greater numbers 
than females. These observations are consistent with other studies which suggest that males precede females to the spawning 
grounds, that females remain there for a short period and that a minimum two males per female is required for reproduction. 
This study shows that the Gatineau River rapids must be protected because they harbor three species that have obtained a status 
by COSEWIC (Threatened: Channel Darter; Special concern: Northern Brook Lamprey and River Redhorse) as well as being 
an important multi-species spawning ground. 


Key Words: Rapids, Gatineau River, Outaouais, spawning, Silver Redhorse, Moxostoma anisurum, River Redhorse, Moxostoma 
carinatum, Shorthead Redhorse, Moxostoma macrolepidotum, Northern Brook Lamprey, Ichthyomyzon fossor, 
Channel Darter, Percina copelandi. 


Un inventaire réalisé au printemps 1989 a l’aide de 
filets de dérive a permis de constater que les rapides 
de la riviére Gatineau en aval du barrage hydroélec- 
trique de Farmer’s Point (Gatineau, Québec) (Figure 1) 
sont utilisés pour la reproduction de I’ Esturgeon jaune, 
des Percidae (Sander spp. et Perca flavescens), et des 
Catostomidae (Catostomus spp. et Moxostoma spp.) 
(Houde et Fournier 1992). Une péche expérimentale 
réalisée en 1998 (données non publiées) a révélé la 
présence a ce site d’une concentration exceptionnelle 
de Chevalier de riviére. Des travaux furent donc entre- 
pris en 1999 pour décrire cette population et son utili- 
sation du rapide Farmer’s. 


L étude ichtyologique de ce secteur de la riviere revét 
un intérét particulier parce que le barrage agit comme 
barriére au déplacement des poissons vers l’amont. Ceci 
pourrait occasionner une concentration de poissons dans 
le secteur des rapides en aval du barrage, en particu- 
lier au printemps, lors des migrations reproductrices. 
Ainsi, le secteur en aval du barrage pourrait étre un site 
privilégié pour la reproduction des poissons provenant 
des riviéres Gatineau et des Outaouais. 

Le but de notre étude est de présenter |’inventaire 
printanier des espéces de poissons de ce secteur de la 
riviére alors que plusieurs de celles-ci sont normale- 
ment en période de fraie. De plus, une attention particu- 


521 


522 


45° 29'21"N 


75° 45'17°O 
t 
po Sz 


AUX 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


45°28'42°N 


75°44'13°O 


outa 


ONTARIO 


Figure 1. Site de la riviere Gatineau prés du pont Alonzo-Wright, se trouvant a 5,2 km de la 


jonction avec la riviére des Outaouais. 


liere sera portée sur la répartition dans le temps de la 
reproduction des trois especes de chevalier (Moxos- 
toma spp.) qui utilisent le secteur des rapides pour 
frayer. 


Site d’échantillonnage 

La section de la riviére Gatineau (45°29'N, 
75°44'O) échantillonnée se trouve entre 6,0 et 5,2 km 
de l’embouchure sur la riviere des Outaouais (Figure 
1). La zone d’échantillonnage couvre une longueur de 
800 m et est délimitée en amont par le pont Alonzo- 
Wright et a l’aval par l’extrémité d’une ile localisée prés 
de la rive ouest de la riviére. La largeur de la riviére 
dans ce secteur varie de 170 m en amont 4 330 m en 


aval et couvre une superficie d’environ 22 ha. La pro- 
fondeur de |’eau varie de 0,5 a 5 m. Le courant est 
particuliérement rapide sur une distance d’environ 
400 m a l’aval du pont et la vitesse devient beaucoup 
plus lente par la suite alors que la riviére s’élargit et 
devient plus profonde. 

Dans les eaux rapides, le substrat est composé de 
galets, de cailloux et de gros blocs épars dans le centre 
de la riviére et de gravier en rive ouest. La rive est a 
été largement modifiée par des remblayages et la 
construction de quais et murs de soutenement. En 
aval, ot la riviére est plus large et le courant plus 
lent, le fond est constitué de gravier, de cailloux et de 
galets; en rive ouest, le substrat est composé de sable 


2004 CoMTOIS ET AL. : L- ICHTYOFAUNE DE LA RIVIERE GATINEAU 523 
TABLEAU I. Espéces recensées lors de l’échantillonnage du rapide Farmer’s, riviére Gatineau, au printemps 1999. 
Famille Nom latin (Espeéce) Nom commun frangais Nombre 
d’individus 
Petromyzontidae Ichthyomyzon unicuspis Lamproie argentée 8 
Ichthyomyzon fossor Lamproie du nord | 
Lampetra appendix Lamproie de I’est 3] 
Acipenseridae Acipenser fulvescens Esturgeon jaune 19 
Hiodontidae Hiodon tergisus Laquaiche argentée 276 
Anguillidae Anguilla rostrata Anguille d’ Amérique | 
Cyprinidae Luxilus cornutus Méné a nageoires rouges 4 
Notemigonus crysoleucas Méné jaune D) 
Notropis atherinoides Méné émeraude 309 
Notropis hudsonius Queue a tache noire 2 
Rhinichthys cataractae Naseux des rapides | 
Semotilus corporalis Ouitouche 52 
Catostomidae Carpiodes cyprinus Couette 68 
Catostomus commersoni Meunier noir 33 
Moxostoma anisurum Chevalier blanc 223 
Moxostoma carinatum Chevalier de riviere 270 
Moxostoma macrolepidotum Chevalier rouge 254 
Ictaluridae Ameiurus nebulosus Barbotte brune 9 
Ictalurus punctatus Barbue de riviére ay 
Noturus insignis Chat-fou liséré 3 
Esocidae Esox lucius Grand brochet l 
Esox masquinongy Maskinongé 4 
Salmonidae Salmo trutta Truite brune | 
Salvelinus namaycush Touladi l 
Percopsidae Percopsis omiscomaycus Omisco 5 
Centrarchidae Ambloplites rupestris Crapet de roche 116 
Lepomis gibbosus Crapet-soleil 9 
Lepomis macrochirus Crapet arlequin 4 
Micropterus dolomieu Achigan a petite bouche 17 
Micropterus salmoides Achigan a grande bouche 1 
Pomoxis nigromaculatus Marigane noire 22 
Percidae Etheostoma nigrum Raseux-de-terre noir 1 
Etheostoma olmstedi Raseux-de-terre gris 12 
Perca flavescens Perchaude 79 
Percina caprodes Fouille-roche zébré 315 
Percina copelandi Fouille-roche gris 76 
Sander canadensis Doré noir 16 
Sander vitreus Doré jaune 80 
Sciaenidae Aplodinotus grunniens Malachigan 5 


et de limon alors que la rive est a aussi été largement 
modifiée par les activités humaines. 


Méthodes 

Léchantillonnage s’est effectué a intervalle de 1 a 
5 jours, du 29 avril au 23 juin 1999, entre 19:00 h et 
01:00 h. Il s’est fait a aide d’un bateau de péche élec- 
trique de marque Smith Root 16R. Des voltages de 
250 a 1000 V et des ampérages de 0,25 a 3,00 A ont 
été utilisés selon la conductivité de l'eau. La péche 
débutait a 20 m des piliers du pont Alonzo-Wright. 
Une dérive contrélée du bateau était faite sur des tran- 
sects de 400 a 800 m le long des rives et au centre de 
la riviere. La période de péche active (lorsque le courant 
était émis dans |’ eau) variait de 320 4 1732 s. Les pois- 
sons capturés étaient déposés dans un vivier installé sur 
le pont du bateau. Une fois le vivier rempli, la péche 
active était arrétée et les poissons étaient examinés et 


relachés, sauf pour quelques spécimens qui ont été 
conservés pour des études ultérieures. 

Les poissons étaient identifiés, dénombrés et le sexe 
ainsi que le stade de reproduction étaient déterminés 
lorsque cela était possible. Pour les chevaliers, le sexe 
était établi a partir de la présence de tubercules nuptiaux 
(Scott et Crossman 1974). Le stade de reproduction 
était déterminé par des caractéristiques morphologiques 
externes et selon l’index de maturité des gonades de 
Nikolsky (1963). Celui-ci classe les stades en six 
catégories. Les stades I, II et Ill, ne sont pas 
identifiables puisqu ils refletent des états de maturation 
des gonades qui ne montrent pas de signes externes. 
Le stade IV est caractérisé par l’identification de 
gonades fermes et gonflées au toucher de la paroi 
abdominale sans expulsion d’ceufs ou de sperme. Le 
stade V est identifié lorsqu’il y a expulsion de produits 
sexuels par simple pression de l’abdomen. 


524 


20 


— 
on 


Nombre d'espéces capturées 
3 


THE CANADIAN FIELD-NATURALIST 


0 0,0 


Vol. 118 


=i 
Ol 
° 


= 
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oO 


oa 
fo) 


29-avr 3-mai 6-mai 10-mai 13-mai 17-mai 20-mai 25-mai 27-mai 31-mai 3-juin 7-juin 10-juin 14-juin 16-juin 17-juin 21-juin 23-juin 


Date 


FicuRE 2. Evolution temporelle du nombre d’espéces capturées et de la température de l’eau (m) au rapide Farmer’s, 


printemps 1999. 


Un poisson au stade VI a un abdomen relativement 
flasque et sans expulsion massive de produits sexuels 
par pression abdominale. 

L’échantillonnage ciblant surtout les chevaliers, |’a- 
bondance des espéces de petite taille est probablement 
une sous-estimation de leur présence dans le milieu. 


Résultats 
Familles et espéces capturées 

L’échantillonnage des rapides de la riviére Gatineau 
a permis de capturer 2388 poissons en 12 heures de 
péche active réparties sur 18 jours entre le 29 avril et 
le 23 juin 1999. Les captures se répartissent en 13 fa- 
milles et 39 espéces (Tableau 1). Les Percidae dominent 
pour le nombre d’espéces capturées (sept espéces). 
Les Cyprinidae et les Centrarchidae sont représentés 
par six espéces chacune. Les Catostomidae dominent 
pour le nombre de spécimens (848), les chevaliers 
constituant 88% des captures. Un seul spécimen des 
Anguillidae (Anguilla rostrata) a été capturé (Tableau 
1). D’ailleurs, un seul spécimen a été capturé pour huit 
especes (Tableau 1) alors que plus de 300 spécimens 
ont été capturés pour le Méné émeraude (309) et le 
Fouille-roche zébré (315). 

Six des espéces capturées ont fait l’objet d’une 
évaluation par le Comité sur la situation des espéces 
en péril au Canada (COSEPAC 2003) : l’Esturgeon 
jaune (non en péril; 1986), le Chevalier de riviére (situ- 
ation préoccupante; 1987), le Chat-fou liséré (menacé; 
1989 et données insuffisantes lors de la révision de 
2002), la Lamproie du nord (situation préoccupante; 
1991), le Fouille-roche gris (menacé; 1993, 2002) et 
le Raseux-de-terre gris (non en péril; 1993). Deux de 
ces especes, le Chevalier de riviére et le Fouille-roche 


gris ont été capturés en grand nombre dans la riviere 
Gatineau, soit 270 et 76 individus respectivement 
(Tableau 1). 

Comme nous pouvons le constater a la Figure 2, le 
plus petit nombre d’espéces capturées a été de dix 
lors de la premiere sortie, le 29 avril. Il a augmenté 
graduellement pour atteindre un maximum de 21 es- 
peces le 27 mai. Par la suite, le nombre a diminué, vari- 
ant de 12 a 17 espéces par sortie. 


Reproduction 

Des individus de 16 des espéces capturées étaient 
au stade V du cycle de reproduction et semblent donc 
y avoir frayé (Tableau 2). Pour plusieurs de ces espéces, 
nos captures permettent de délimiter une période de 
reproduction probable et les températures auxquelles 
elle se serait produite. Pour quelques espéces, des in- 
dividus en état de fraie (stade V) étaient présents dés 
la premiere journée d’échantillonnage ou encore pré- 
sents la derniére journée. Dans ces cas, il est possible 
que la période de reproduction s’étende au-dela de la 
période d’échantillonnage. Des individus des deux 
sexes de tous les stades avancés de développement des 
gonades (stades IV, V, VI) ont été observés pour cing 
especes seulement : le Chevalier blanc (Moxostoma 
anisurum), le Chevalier de riviere (M. carinatum), le 
Chevalier rouge (M. macrolepidotum), la Couette (Car- 
piodes cyprinus) et la Laquaiche argentée (Hiodon 
tergisus). 


Reproduction des trois espéces de chevalier 
L’abondance des captures permet d’établir la chro- 
noséquence des périodes de reproduction des trois 
especes de chevaliers dans la riviére Gatineau (Figures 
3 et 4). Pour toutes les espéces, des spécimens de 


2004 


COMTOIS ET AL. : L;: ICHTYOFAUNE DE LA RIVIERE GATINEAU 


Nn 
i) 
Nn 


TABLEAU 2. Période de capture d’individus en fraie (stade V), selon le sexe et ’espece, au rapide Farmer’s, printemps 1999. 


Espéce Nombre d’individus 
fe) Q 
Ambloplites rupestris 2 3 
Aplodinotus grunniens | | 
Carpiodes cyprinus 35 7 
Catostomus commersoni 8 5 
Hiodon tergisus 198 2 
Ichthyomyzon fossor 0 | 
Ichthyomyzon unicuspis 2 
Lampetra appendix 5 3 
Moxostoma anisurum 89 6 
Moxostoma carinatum 96 16 
Moxostoma macrolepidotum 67 7 
Percina caprodes 10 0 
Percina copelandi d/ 1 
Percopsis omiscomaycus 0 2 
Pomoxis nigromaculatus 3 3 
1 1 


Période de capture Température de 


eau (°C)! 

27 mai — 10 juin 13,5 — 18,5 

3— 6 mai 9,0 — 10,0 

29 avril — 3 juin 7,0 — 18,0 

29 avril — 27 mai 7,0 — 13,5 

10 mai — 23 juin 12,5 — 19,0 
23 juin 19,0 

13 mai — 23 juin 12,5 — 19,0 

3 — 20 mai 9,0 — 14,0 

3 — 31 mai 9,0 — 17,0 

10 mai — 23 juin 12,5 — 19,0 

29 avril — 21 juin 7,0 — 19,0 

6 — 25 mai 10,0 — 14,0 

20 mai — 21 juin 14,0 — 19,0 
6 mai 10,0 

3 — 13 mai 9,0 — 12,5 

3 — 20 mai 9,0 — 14,0 


Semotilus corporalis 


' Température de l’eau au début et a la fin de la période de capture. 


stade de maturité IV étaient présents au début de la 
période d’échantillonnage, le 29 avril, sauf pour les 
femelles du Chevalier rouge et les males du Cheva- 
lier de riviere. Pour ceux-ci, les individus au stade [V 
sont apparus le 3 et le 6 mai respectivement (Figure 
3). Des males de stade IV ont été capturés jusqu’au 


17 mai dans le cas du Chevalier rouge, jusqu’au 27 
mai pour le Chevalier blanc et jusqu’au 7 juin pour le 
Chevalier de riviere. Les dernieres femelles de stade 
IV de Chevaliers rouge et blanc ont été capturées le 
17 mai, alors qu’elles ont été présentes sur le site 
jusqu’au 17 juin pour le Chevalier de riviere. 


(8 enna 
(34) 

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0 i ee ey 
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ae 

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= 3 (96) —— ee Deuces 
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3 Ca NER SE CREE RUT SEL Se a Le Ee 2) 

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| 
f - + - } — + —— — : oo 
26-avr 1-mai 6-mai 11-mai 16-mai 21-mai 26-mai 31-mai 5-juin 10-juin 15-juin 20-juin 25-juin 
Date 


FIGURE 3. Période de reproduction des trois espéces de chevalier selon l’index de maturité de Nikolsky (1963) dans la riviére 
Gatineau, au printemps 1999. Le carré représente le Chevalier blanc (Mi), le losange, le Chevalier de riviére (@) et le 
triangle, le Chevalier rouge (a). Les symboles pleins désignent les femelles tandis que les vides représentent les males. 
Le nombre d’individus capturés est indiqué entre parenthéses. Les lignes verticales correspondent au début et a la fin 
de la période d’échantillonnage. Le pointillé 4 l’extérieur de la période d’échantillonnage signifie que des individus 
appartenant aux stades indiqués ont pu se retrouver sur la frayére avant ou apres cette période. 


526 


Des males au stade V sont apparus dans les cap- 
tures le 3 mai pour le Chevalier blanc et le 10 mai 
pour le Chevalier de riviére, 4 une température de 9,0 
et 12,5 °C respectivement. Les males Chevalier rouge 
de ce stade de maturité étaient présents des le début de 
l’échantillonnage. Les femelles de stade V sont appa- 
rues le 10 mai pour le Chevalier blanc et le Chevalier 
rouge (12,5 °C) et le 3 juin pour le Chevalier de rivi- 
ére (18,0 °C). Les captures de males au stade V sont 
toujours nettement plus abondantes que celles des fe- 
melles. Aucune femelle de stade V n’a été capturée 
aprés le 20 mai dans le cas du Chevalier blanc (14,0 °C) 
et le 7 juin dans le cas du Chevalier rouge (18,5 °C) 
alors qu’elles étaient toujours présentes a la fin de I’é- 
chantillonnage, le 23 juin, dans le cas du Chevalier de 
riviere, 4 une température de 19,0 °C. 

Des femelles au stade VI apparaissent dans les cap- 
tures le 17 mai pour le Chevalier blanc et le Chevalier 
rouge (12,5 °C) alors qu’elles sont présentes pour le 
Chevalier de riviére 4 compter du 10 juin (18,5 °C). 
Les dernieres femelles de stade VI ont été capturées 
le 31 mai pour le Chevalier blanc (17,0 °C) et le 17 juin 
pour le Chevalier rouge (19,0 °C), alors qu’elles étaient 
toujours présentes a la derniere date d’échantillonnage 
pour le Chevalier de riviere (19,0 °C). 


Discussion 

Si l’on ajoute aux 16 espéces citées dans le Tableau 
2 celles rapportées par Houde et Fournier (1992), il est 
probable qu’environ 20 espéces se reproduisent dans le 
rapide Farmer’s. D’autres espéces capturées pourraient 
potentiellement utiliser le secteur échantillonné de la 
riviere Gatineau pour frayer (Scott et Crossman 1974; 
Bernatchez et Giroux 2000) : le Méné émeraude, le 
Naseux des rapides, et possiblement le Méné a nage- 
oires rouges et le Raseux-de-terre gris. Toutefois, la 
période d’échantillonnage ne correspondait pas a la 
période de fraie pour certaines espéces (e.g. Méné 
émeraude) alors que trop peu de spécimens ont été cap- 
turés pour d’autres afin de déterminer |’ état reproduc- 
tif (e.g. Naseux des rapides). 

La frayére multispécifique des rapides de la riviére 
Gatineau compte autant d’espéces que celle de la 
riviere aux Pins, a Boucherville (Massé et al. 1988). 
Les deux frayeres ont toutefois des caractéristiques trés 
différentes; la frayére de la riviére aux Pins constitue 
un habitat d’eau calme utilisé particuliérement par 
les espéces phytophyles. D’ailleurs, seulement trois 
especes sont communes aux deux frayéres : la Oui- 
touche, le Meunier noir et le Crapet de roche. Dans 
les eaux vives, Massé et al. (1988) mentionnent cing 
autres frayéres entre Carillon (riviére des Outaouais) 
et la riviere des Prairies qui comptent entre 8 et 27 
especes reproductrices. La riviére Gatineau se situe 
donc dans le milieu du peloton quant a son utilisation 
multispécifique. 

Des femelles en état de fraie de Chevaliers rouge et 
blanc sont apparues sur le site en méme temps le 10 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


mai. Elles y ont été capturées pendant une période de 
10 jours dans le cas du Chevalier blanc et de 28 jours 
dans le cas du Chevalier rouge. La fraie des deux es- 
peces a donc été en partie simultanée (Figures 3 et 4). 

Contrairement a nos résultats, Curry et Spacie (1984) 
ont noté que le Chevalier blanc a amorcé la reproduc- 
tion une semaine avant le Chevalier rouge dans le Deer 
Creek, Indiana, en 1979. Toutefois, Mongeau et al. 
(1986, 1992) ont également observé une simultanéité 
de la reproduction du Chevalier blanc et du Chevalier 
rouge dans leur étude sur la riviere Richelieu. En effet, 
ils ont observé que la reproduction de ces deux espéces 
débutait 4 la mi-mai pour se terminer a la mi-juin. Nos 
données semblent tendre vers le méme résultat puisque 
des individus de stade VI ont été capturés du 17 mai 
au 21 juin pour les deux espéces (Figure 3). Mongeau 
et al. (1986, 1992) ont noté que le Chevalier rouge et 
le Chevalier blanc commengaient a déposer des ceufs 
a une température de 10,7 °C. Meyer (1962) a indiqué 
que la température a laquelle la reproduction débutait 
pour le Chevalier rouge et le Chevalier blanc sur la 
riviere Des Moines, en Iowa, était 11,1°C et 13,3°C 
respectivement. Par contre, Burr et Morris (1977) ont 
observé une température de 16,0°C pour la reproduc- 
tion du Chevalier rouge dans le ruisseau Big Rock, 
en Illinois et Hackney et al. (1970), une température 
de 14,4°C pour le Chevalier blanc dans la riviere Flint, 
en Alabama. La Figure 2 montre que la température 
de l’eau au 17 mai était a 12,5°C, ce qui correspond 
relativement bien aux données de Meyer (1962) et de 
Mongeau et al. (1986, 1992). 

Burr et Morris (1977) indiquent que plusieurs es- 
peces de chevaliers pourraient possiblement frayer a 
des périodes et des températures similaires et sur des 
substrats comparables. II peut toutefois y avoir de la 
ségrégation spatiale entre les espéces. En effet, le Che- 
valier rouge et le Chevalier doré (Moxostoma erythru- 
rum) ont été observés en train de frayer simultanément 
au méme site, mais le Chevalier doré avait une dis- 
tribution plus étendue en amont, ce qui lui fournissait 
un habitat non utilisé par le Chevalier rouge (Curry et 
Spacie 1984). Par ailleurs, une étude comparant la fraie 
chez le Chevalier doré et le Chevalier noir (Moxostoma 
duquesnei) a démontré que ces deux espéces se repro- 
duisaient au méme moment et dans la méme section 
du Stony Creek, en Illinois (Kwak et Skelly 1992). 
Les auteurs mentionnent toutefois qu’il existe des dif- 
férences dans le choix des microhabitats de fraie et dans 
le comportement reproducteur et que celles-ci servi- 
raient possiblement a réduire la compétition entre ces 
espeéces. Bien qu’une étude plus approfondie de l’habi- 
tat et du comportement reproductif soit nécessaire, il 
est possible qu’une ségrégation spatiale et/ou compor- 
tementale se produise pour le Chevalier blanc et le 
Chevalier rouge dans la riviére Gatineau. 

Des femelles de stade V du Chevalier de rivieére ont 
été capturées pour la premiere fois le 3 juin et ce, 
jusqu’a la derniére journée d’échantillonnage. Sa re- 


2004. COMTOIS ET AL. : L;- ICHTYOFAUNE DE LA RIVIERE GATINEAU ayy 


35 + 


Chevalier blanc 
| Femelle: n= 51 
30 7 Male: n= 94 
25 Individus non sexés: n= 78 


— 
nn 
a 


i Femelle 
[_] Male 


Ue eee 


25 7 Chevalier de riviére 


Femelle: n = 110 
Male: n = 133 


w” 

@ 

ie 

= 20 

e | Individus non sexés: n = 27 

a) j 

3 15 

2 

GT = 

= 1055; 

5 i 

§ 

£ Sy 

6 ; 

Zz ] 
0 


20 7 Chevalier rouge 


Femelle: n= 70 
Male: n= 108 
15 -  Individus non sexés: n= 76 


| 1 


10 


| ae 


Oo on 
cs eee 
peipiccos nace 
DS rae are ae) 
| Sea 
= 

i 

i 
ke 

juin 


[Cl |e ee] 


a Ln lan 
=> OCG G cs eS - SS SS Ss = 
gee ee eo Ssocas 
is 
A Se SAAN: SAN 


FiGurE 4. Nombre d’individus capturés par jour selon le sexe pour les trois espéces de chevalier, rivi¢re Gatineau, printemps, 


1999. 


528 


production débute donc plus tard, soit autour du 10 
juin (Figure 3) a une température de 18,5 °C. Mongeau 
et al. (1986, 1992) ont obtenu plus ou moins les mémes 
résultats puisqu’ils indiquent que le Chevalier de riviere 
amorce la fraie autour de Ja deuxi¢me semaine de juin 
a une température de 17,7 °C, une conclusion fondée 
sur la capture de deux spécimens au stade VI. Hackney 
et al. (1970) mentionnent également que le Chevalier de 
riviére a une reproduction tardive vis-a-vis de celle 
du Chevalier blanc dans la riviére Duck, au Tennessee. 
La différence entre la période de reproduction de cette 
espece et celles des deux autres chevaliers présents 
dans les rapides de la riviére Gatineau est probablement 
due au fait que le Chevalier de riviére nécessite une 
température de |’eau plus élevée pour débuter sa 
reproduction. 

Par ailleurs, pour ces trois espéces, nous observons 
des males de stade V a une date plus hative que les 
femelles au méme stade (Figure 3). Ceci s’explique par le 
fait que les males du Chevalier blanc, du Chevalier rouge 
et du Chevalier de riviere précédent les femelles sur 
les frayeres (Meyer 1962; Hackney et al. 1968, 1970; 
Scott et Crossman 1974). Ce phénoméne est d’ailleurs 
observé chez d’autres chevaliers et chez plusieurs 
autres Catostomidae (Meyer 1962; Scott et Crossman 
1974; Jenkins et Jenkins 1980; Page et Johnston 1990; 
Kwak et Skelly 1992; Cooke et Bunt 1999), 

Dans nos captures, le nombre de males de stade V est 
plus grand que celui de femelles pour les trois espéces 
de chevalier. Ceci n’est pas inhabituel. Hackney et al. 
(1970) ont observé un rapport des sexes de 4 males : | 
femelle sur une frayére de Chevalier blanc. De plus, 
Meyer (1962) note que chez les Chevaliers rouge et 
blanc, les femelles demeurent sur les sites de fraie 
beaucoup moins longtemps que les males. La fraie chez 
les chevaliers, et chez la plupart des Catostomidae, se 
fait généralement en présence de deux males et d’une 
femelle (Hackney et al. 1968; Burr et Morris 1977; 
Jenkins et Jenkins 1980; Page et Johnston 1990; Kwak 
et Skelly 1992; Cooke et Bunt 1999), d’ot la nécessité 
d’une présence en plus grand nombre de males sur la 
frayére tel que nous l’avons observé pour chacune des 
trois espéces présentes au rapide Farmer’s. 

Nos travaux ont permis de découvrir le site de re- 
production de l’une des plus importantes populations 
de Chevalier de riviére. Un échantillonnage sur quel- 
ques années serait nécessaire pour préciser la chrono- 
séquence de fraie de cette espéce et des autres che- 
valiers utilisant le site. Un échantillonnage sur une 
plus longue période aurait probablement pu permettre 
de confirmer la fraie au rapide Farmer’s d’un plus grand 
nombre d’espéces. 

Cette étude démontre donc que le rapide Farmer’s 
constitue une frayére multispécifique importante tant 
par le nombre d’espéces qui s’y reproduisent que par la 
présence d’espéces évaluées par le COSEPAC comme 
ayant une situation préoccupante (Lamproie du nord, 
Chevalier de riviére) ou méme menacée (Fouille-roche 
gris). Localisés en plein coeur d’une zone urbaine, ces 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


rapides ont une grande valeur écologique et doivent 
étre protégés. 


Remerciements 

Nous remercions Anne Phelps, Annie-Chantal Gui- 
bord et Kevin Moon pour leur aide sur le terrain. Cette 
recherche a été partiellement subventionnée par une 
subvention a la découverte du Conseil de recherches en 
sciences naturelles et en genie du Canada (CRSNG) 
[Frangois Chapleau]. 


Littérature Citée 

Bernatchez, L., et M. Giroux. 2000. Les poissons d’eau 
douce du Québec et, leur répartition dans l’Est du 
Canada. Broquet, Boucherville, Québec. 350 pages. 

Burr, B. M., et M. A. Morris. 1977. Spawning behavior of the 
Shorthead Redhorse, Moxostoma macrolepidotum, in Big 
Rock Creek, Illinois. Transactions of the American Fish- 
eries Society 106: 80-82. 

Cooke, S. J., et C. M. Bunt. 1999. Spawning and reproduc- 
tive biology of the Greater Redhorse, Moxostoma valen- 
ciennesi, in the Grand River, Ontario. Canadian Field- 
Naturalist 113: 497-502. 

COSEPAC. 2003. Résultats des évaluations du COSEPAC, 
novembre 2003. Comité sur la situation des espéces en 
péril au Canada, Ottawa, Ontario. 54 pages. 

Curry, K. D., et A. Spacie. 1984. Differential use of stream 
habitat by spawning catostomids. American Midland Nat- 
uralist 111: 267-279. 

Hackney, P.A., W.M. Tatum, et S.L. Spencer. 1968. Life his- 
tory study of the River Redhorse, Moxostoma carinatum 
(Cope), in the Cahaba River, Alabama, with notes on the 
management of the species as a sport fish. Proceedings of 
the Annual Conference of the Southeastern Association 
of Game and Fish Commissioners 21: 324-332. 

Hackney, P. A., G. R. Hooper, et J. F. Webb. 1970. Spawn- 
ing behavior, age and growth, and sport fishery for the 
Silver Redhorse, Moxostoma anisurum (Rafinesque), in the 
Flint River, Alabama. Proceedings of the Twenty-Fourth 
Annual Conference, Southeastern Association of Game 
and Fish Commissioners 1970: 569-576. 

Houde, P. et H. Fournier. 1992. Travaux de recherche de 
frayéres dans la riviére des Outaouais et ses principaux 
affluents au cours du printemps 1989. Ministére du Loisir, 
de la Chasse et de la Péche, Service de l’aménagement et 
de l’exploitation de la faune, Hull. 28 pages. 

Jenkins, R. E., et D. J. Jenkins. 1980. Reproductive behavior 
of the Greater Redhorse, Moxostoma valenciennesi, in the 
Thousand Islands region. Canadian Field-Naturalist 94: 
426-430. 

Kwak, T. J., et T. M. Skelly. 1992. Spawning habitat, behav- 
ior, and morphology as isolating mechanisms of the Gol- 
den Redhorse, Moxostoma erythrurum, and the Black Red- 
horse, M. duquesnei, two syntopic fishes. Environmental 
Biology of Fishes 34: 127-137. 

Massé, G., R. Fortin, P. Dumont, et J. Ferraris. 1988. Etude 
et aménagement de la frayere multispécifique de la rivi- 
ére aux Pins et dynamique de la population de Grand bro- 
chet, Esox lucius L., du fleuve Saint-Laurent, Boucher- 
ville, Québec. Ministére du Loisir, de la Chasse et de la 
Péche, Service de l’aménagement et de l’exploitation de 
la faune, Montréal, Québec, Rapport technique 06-40. 
224 pages. 


2004 


Meyer, W. H. 1962. Life history of three species of redhorse 
(Moxostoma) in the Des Moines River, Iowa. Transactions 
of the American Fisheries Society 91: 412-419. 

Mongeau, J.-R., P. Dumont, et L. Cloutier. 1986. La biolo- 
gie du Suceur cuivré, Moxostoma hubbsi, une espeéce rare 
et endémique a la région de Montréal, Québec, Canada. 
Ministére du Loisir, de la Chasse et de la Péche, Service 
de l’aménagement et de l’exploitation de la faune, 
Montréal, Québec, Rapport technique 06-39. 137 pages. 

Mongeau, J.-R., P. Dumont, et L. Cloutier. 1992. La biolo- 
gie du Suceur cuivré (Moxostoma hubbsi) comparée a celle 
de quatre autres espéces de Moxostoma (M. anisurum, M. 
carinatum, M. macrolepidotum, M. valenciennesi). Revue 
canadienne de zoologie 70: 1354-1363. 


COMTOIS ET AL. : L- ICHTYOFAUNE DE LA RIVIERE GATINEAU 


529 


Nikolsky, G. V. 1963. The ecology of fishes. Traduit par L. 
Birkett. Academic Press, New York. 352 pages. 

Page, L. M., et C. E. Johnston. 1990. Spawning in the Creek 
Chubsucker, Erimyzon oblongus, with a review of spawn- 
ing behavior in suckers (Catostomidae). Environmental 
Biology of Fishes 27: 265-272. 

Scott, W. B., et E. J. Crossman. 1974. Poissons d’eau douce 
du Canada. Office des recherches sur les pécheries du 
Canada Bulletin 184. 1026 pages. 


Received 3 October 2003 
Accepted 21 June 2004 


Moths and Butterflies (Lepidoptera) of the Boreal Mixedwood Forest 
near Lac La Biche, Alberta, Including New Provincial Records 


Grea R. Pout!, Davip W. LANGor!, JEAN-FRANCOIS LANDRY’, and JOHN R. SPENCE? 


' Natural Resources Canada, Canadian Forest Service, 5320 - 122 St., Edmonton, Alberta T6H 3S5 Canada 

> Agriculture and Agrifood Canada, Eastern Cereal and Oilseed Research Centre, 960 Carling Avenue, Ottawa, Ontario 
K1A 0C6 Canada 

> Department of Renewable Resources, University of Alberta, Edmonton, Alberta T6G 2H1 Canada 


Pohl, Greg R., David W. Langor, Jean-Francois Landry, and John R. Spence. 2004. Moths and Butterflies (Lepidoptera) of 
the boreal mixedwood forest near Lac La Biche, Alberta, including new provincial records. Canadian Field-Naturalist 
118(4): 530-549. 


Lepidoptera were collected, primarily via UV light trap, for three seasons in the boreal mixedwood forest near Lac La 
Biche, Alberta. A total of 11 111 specimens were collected, representing 41 families and 438 species. A species list with 
flight times is presented. The total Lepidoptera community was estimated to be 546 +23.34 species. Abundance and species 
richness peaked in late July. Thirty-five species constitute new records for Alberta, while one species, Acanthopteroctetes 
bimaculata, is a new record for Canada, and the first record of the family Acanthopteroctetidae in Canada. 


Key Words: Moths, butterflies, Lepidoptera, Lac La Biche, Alberta, flight times, Acanthopteroctetes bimaculata, new to 


Canada. 


Alberta is blessed with an abundance and variety 
of forested lands. A large portion of the province is 
covered by the boreal mixedwood (Figure |) domi- 
nated by Trembling Aspen (Populus tremuloides), 
Balsam Poplar (P. balsamifera), and White Spruce 
(Picea glauca) and containing lesser amounts of White 
Birch (Betula papyrifera), Balsam Fir (Abies bal- 
samea), and other species (Beckingham and Archibald 
1996). Over the last 15 years Trembling Aspen has 
increased greatly in value as a commercial tree species 
and there has been a large increase in forestry activity 
centered on this resource. A large portion of Alberta’s 
aspen mixedwood forest is now scheduled for har- 
vesting over the next 30-40 years (Pratt and Urquhart 
1994). It is a major concern that our knowledge of these 
forests is relatively poor and there is little empirical 
basis for predicting the impacts of harvesting and other 
development on non-timber values such as biodiversity. 

In an effort to obtain baseline data for assessing 
impacts of forestry practices on biodiversity, and to 
determine whether old aspen stands in mixedwood 
forests contain unique species, a multi-agency team 
of scientists studied the structure and composition of 
biotic assemblages in aspen-dominated forests of 
various ages in the vicinity of Lac La Biche, Alberta, 
between 1993 and 1995. This work is among the most 
comprehensive biotic inventories in aspen forests in 
the province. Lists of plants, birds, amphibians, and 
mammals have been published by Stelfox (1995) and 
those of ground-dwelling beetles and dead wood- 
inhabiting beetles by Spence et al. (1997) and Ham- 
mond (1997), respectively. As part of that study, but- 


terflies and moths were also sampled. This represents 
the first concerted effort to inventory Lepidoptera, 
especially moths, in aspen forests in western Canada. 
Lepidoptera constitute a major component of boreal 
forest biodiversity (Danks and Foottit 1989), and are 
important herbivores and pollinators (Scoble 1992). 
The abundance of new provincial records among micro- 
moths (defined here as the primitive and monotrysian 
groups, and the lower ditrysian superfamilies up to 
and including the Pyraloidea and Thyridoidea sensu 
Kristensen 1999) indicates how poorly sampled these 
groups have been. The attached checklist and flight 
times provide baseline information for comparison to 
other studies and to aid in future research. 


Materials and Methods 

The study area (Figure 1) is located in Lakeland 
Provincial Park near Touchwood Lake, east of Lac 
La Biche (54°51'N, 111°27'W) in the Central Mixed- 
wood subregion of the Boreal Forest Natural Region 
(Beckingham and Archibald 1996). Lepidoptera were 
sampled in two stands: a 65-year-old (“mature”) stand 
of 269 Ha, containing 83% cover of Trembling Aspen, 
15% Balsam Poplar, 2% willow (Salix spp.), and 1% 
White Birch, and a stand over 130 years old (“old”) 
of 148 Ha, containing 54% cover of Trembling Aspen, 
32% White Birch, 11% Balsam Poplar, and 3% willow. 
An inventory of vascular plants found around these 
study sites is included in the lists published by Stelfox 
(1995). Both stands were of fire origin, and were largely 
undisturbed by humans. The mature stand is consid- 
ered to be of rotation age and the old stand is much 


530 


2004. POHL, LANGOR, LANDRY, and SPENCE: 
‘ ae ies - : e 
cat Edmonton waa 
ALBERTA 


100 km 


boreal 
mixedwood 


Ficure |. Extent of the boreal mixed wood ecoregion in 
Alberta and location of the study area. 


older than the planned stand rotation age (60-70 years) 
for Alberta aspen forests. A comparison between these 
two stands, based on the light trap catches of Lepi- 
doptera described here, has been done elsewhere (Pohl 
et al. 2004). 

Two 30 watt UV traps were run in each stand, from 
dusk to dawn for one night approximately every two 
weeks, from 16 June to 16 September in 1993, 3 May 


TABLE |. Abundance and diversity of three groups of Lepi- 
doptera collected near Touchwood Lake, Alberta. Butterflies 
include the superfamilies Hesperioidea and Papilionoidea; 
macro-moths include the superfamilies Lasiocampoidea, 
Bombycoidea, Drepanoidea, Geometroidea, and Noctuoidea; 
micro-moths comprise all other superfamilies. 


Group Number of Specimens Number of Species 

(Proportion) (Proportion) 
micro-moths 3897 (0350) 201 (0.470) 
macro-moths 7163 (0.643) 229 (0.523) 
butterflies 51 (0.005) 8 (0.018) 
total TOE a GL 438 


LEPIDOPTERA NEAR LAC LA BICHE 


to 5 October in 1994, and 28 May to 28 August in 
1995. Traps were hung at approximately 1.6 m from 
the ground, and activated from dusk until dawn. These 
samples were augmented by periodic hand collecting 
at portable UV lights, and a small amount of net col- 
lecting of day-flying species. Specimens were identi- 
fied using a wide array of taxonomic publications, and 
by comparing to specimens in the Canadian Forest 
Service’s Northern Forestry Centre Research Collec- 
tion (NFRC) in Edmonton, Alberta, and the Canadian 
National Collection (CNC) in Ottawa, Ontario. Vouch- 
er specimens have been deposited at the NFRC and 
CNC. 

To obtain an estimate of the total size of the Lepi- 
doptera community in the study area, a Chao-1 estimate 
of diversity (Chao and Lee 1992) was calculated as 
described by Colwell and Coddington (1994). 


Results 

A total of 11 111 specimens were collected, repre- 
senting 41 families and 438 species (Table 1). A list of 
all species collected appears in Table 2. Some species, 
particularly micro-moths, are identified here merely as 
morphospecies, reflecting the lack of knowledge of 
the group, and the difficulty in making species identi- 
fications. Examples of some of the micro-moth species 
collected appear in Figure 2; some of the macro-moths 
are illustrated in Figures 3 and 4. The Chao-1 calcula- 
tion estimated the total size of the Lepidoptera com- 
munity in the study area to be 546 + 23.34. 

The 1994 data is examined in detail here, since it 
was the most extensively sampled year and there were 
no trap failures. In 1994, both abundance (Figure 5) 
and species richness (Figure 6) peaked dramatically 
in late July. A major flush of Noctuidae occurred in 
late July, with the highest species richness occurring 
throughout July and into early August. There were 
modest peaks of noctuid species that overwinter as 
adults, in early May and mid-September. The Geo- 
metridae and other macro-moths peaked slightly earlier 
than the Noctuidae, exhibiting maximum abundance 
in early July, and the greatest richness from mid-June 
to late July. The abundance of micro-moths peaked in 
late July, although most of this dramatic peak was a 
single species, Scoparia biplagialis, with 846 speci- 
mens. The greatest micro-moth richness occurred 
throughout July. More modest peaks in micro-moth 
abundance and richness occurred in mid-June and 
mid-September. 


Discussion 

This study likely did not collect all the species pres- 
ent in the study area; the Chao-1 estimator suggests 
that approximately 110 species were missed. Many 
of these are undoubtedly species that are not easily 
sampled via light traps. For example, the amount of 
effort spent collecting butterflies and day-flying moths 
was minimal and many of these taxa were likely missed. 


THE CANADIAN FIELD-NATURALIST Vol. 118 


Da 


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THE CANADIAN FIELD-NATURALIST Vol. 118 


534 


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isnsny Aye ~ UOJUILIS pyjajnuspsyd pIDAVUUTT aepisiiajdouso) 
pollog IWSI[q SOION Jloyiny satoads snuan Ayrurey 


‘Apnjs IaXK-JdIY} IY} JQAO VPLU SUOTDIT[OO UO paseg SI poltad JYSIpy “ISIP{IOYS (€Q6T) “[e 19 SASPpOH IY} SMOTIOJ 
Bxv} JOYSIY UTyIM IoUONbes soa1sadg “(666 ) UASUD}SIDY JO JLY] SMOT[OJ UONBOYTSse]O JOYSIP ‘vIOQLY ‘aye poomyonoy Avau paj}oo]][Oo saroads vsajdopide’q Jo ysvq (panuiuod) 7 AAV], 


535 


IRA NEAR LAC LA BICHE 


iPIDOPTI 


NCE: LE 


iOR, LANDRY, and SPE 


POHL, LANG 


2004 


{yn avy — oun piu (Gueltte)ie)) punjoiospf puilouosd&y) 

snsny Ayia — Ayn ae] (WURYSUIS|e A ) DUDUIUJNI DIJAIOJON 

ARIA] OR] Z (ysnouunqop) DUDBAIAUOD DJBUDY 

Ayn ae] © (SUOLU]D) puppulipd vjaupyd 

oune prw G (Oya) DupawamD AIwau saidods DIaUvY_ 

Ayng pru (OJ.vay,) DupayAng DIUNAY 

Ayng piu — ounr aye] (YIOMLH) DUDINUIUIP S1JKOUY 

oung av] (4o1]O.14) punjdwoo syaouy 

Ayng pr — sung ae] Z (SUdUO]D) pupmplooasnf rweau soroods sijdouy 

oun aye] — oun piu TZ (19][97,) DUuvINIIV] WedU Satoads syAouy 

aune aye] — Aepy qe] (19][97,) pupnbapgns syaouy 

oung oye] — dune pr (souqny) DUDIIIDJAU SaNasysa|Q 

isnsny Ayiea — sung pri (J9TYOSO\) DUDIIDIS SaNadyja|Q 

Ayn piu (UIBYSUIS|eAA ) xajdnp vjiydpiosopnas q 

ysnsny Apiva — Ayn aye] (WOJIvIy) pupaouas smuojody 

Ajng pri — sung aye] (USTIUIOH) ppyful snuojody 

ysnsny Ayia (OJIN) puvjdasap siuojody 

ysnsny Ayjieva — Ane Apiea (aouqnH) puvasdps snuojody 
Ayng pruu (OJ.1vay) DUDAUIO]D DAUD. — IVUTJNIAYJOTO — IePIOL.AAO], 

ysnsny ov] — oune pr (Gelttc)ie)) DUDISAMA Sisda]) 

Ajng aie] — oun pru (AD4[VMA) pDuvonapjaut sisda]) 

Ajng ae] (pyeusa) pupisuauia]o sisda]y 

ysnsny Ayivo — sung oye] (you) puvoisiad sisdayy 

AVI AP] (19q[RMA) puvjoyfy snuapuas 

Ayng aye] — Apne piu (TO4[PAA) DIdsosk8AV SAIYMY 

Ayng pr (SUOWI[D) puvsafnlunf{ DANIUOISIAOYD 

Ayn piu — sung 29] (JO4TRMA) DUDJIIJUOI DANAUOJSIAOYD 

Avy 23¥] (ADq[RMA) DUDIUDG]D DANIUOISIAOYD 

ysnony Ayres — A[ne oye] (SLB) DUDAIVDSOA DANIUOJSIAOY, 

ysnsny Apia — Ang pr NOsAVIyY DUDPVUDI SuUaPUD 

ysnony ae — Apne pr (ADYTVMA) SIDSnavp! VIOUAID] J 

Ajng 33e] | /Satoeds sryjouns.ivdg 

ysnsny Ajiea (SUSU9]-)) DUVIDINIVas Siyjouvssvdsy 

ysn3ny Ajiea (JOY[PAMA) saployjupx s1yjouvnssvds 

oun pru (snovuurT) DUDAJSIUMU DIN 

qsnsny Aiea — A[ng oye] ig (UOSUIqOY) puvjdwuosd sayjay 

oune 3je] (YIOMe}) puvu syayIo) 

Jaquiaydag piu (snto1ige4) DUDSADULA SIAAJIV 

oung Aiea — Jsnsny ae] (uosuIqoYy) DAUIJOABIU SIAAJIV 
poled IsIf4 Sd1ION JouNYy soroads snuay Ayre] 


‘Apnys IWIK-IdIY} OY} IOAO IpRUL SUOT}DIT[OD UO pase SI POLlod IYSIP “ISIPJOOYS (E86) ‘[e 19 SOSPpOH IY} SMOT[OJ 


Bxe] JOYSIY UTYIM BdUANbas sa1dadg “(GHG ) UBSUAISTLY JO VY) SMO[[OJ UONROYISSe]O JOYSIP{ “eLIOq[Yy ‘aye poomyYonoL, Iau pojoa[joo saroeds vsaydopidey Jo isi] (panuyuod) *Z ATAV |, 


Vol. 118 


THE CANADIAN FIELD-NATURALIST 


536 


Ayn ae] (sudua]a) DIJA8VAI3 NA YjYdlss3y 
Ajng oe] (13 [9Z) DIJAJOIANA DALY AUSY 
snsny Ayre p (udyourz) SN]JAYIVA] SNQUIDAD 
snsny oie] — Apne oye] (1jodo9s) snpjajsad snquivsg 
ysnsny Ajaewo — A[ng prut (aehq) SIDj4aqV viuopny 

ysnsny avy — Apne prut IOC sypisvidig viapdoos aepriquivig 
ysndny Ayes — sung aye] (19][97Z) DIJAIUOAPIAYIO VISOINT 
sung Ajiwa — Avy Ay.eo (douqnH ) DIJA1AVINSSO1s DIpoydoz 
ysnony Apres — sung aye] dolUNY 2 BNNINIAY Saplopjanauad DIAJICAOIG 
ysnony Aiwa — Avyy Oe] (jouosey) Dyjaoiajajqns sisdojakpy 
Ajng aye] G \S[NH DiJajniag ‘qoid ‘ds sispqgoioy 

Ayng pruu (1OYIPAA) SIpsnjaucy] DIUOYINOG oeprypeskg 

Avy Ape (UIeYsuUIs|eAA ) DIJajuagSa1d DIpUuOg oeprursodieg 
Avy Ape (WIBYSUIS|eAA ) void pyyddjquy 

Ayng pri (AD4[VAA) SnjAJavpouloy VisUul]|ay] aeploydosaid 

Av] prul Alpue’] 29 Arpuey louuryey Buon] y oepnionyy 

oun ae] — sung Ayre vd (puenig) Dyjajoundi.sadsp V1IyIOM depIpoly) 
Ajng ae] (1ouqnH) DuvIp S1jnasoyy 

oune pr (aekq) DJauaplgIgo SVALO]DI aepynai0yg 
Ayng pr (YOLIUIoH ) pnbopixayf vps) 
Ayn ajey — Avyy aie] (josng) puvjndod pips) 
Avy, oie] — Avy Ape WUBYSUISTeAA pupjoun] DyoydviH, 
Joquiaydag piw — jsnsny ary] (peut) puppul] pyouldy 
ysnsny ary — Apne aye] Z (Hoyeay) pupuowou pyouldy 
Ajng prtu — oun oye] (1OYIPAA) DUDSsusUudA] DYOUIAT 
Jaquiaydag piu (ojIvay) pubajppi4o pyouldy 
Jaquiajydag piu — Avy aye] (o19]D) pijasiu pyouldy 
Ayn avy — Avyy avy] (WIeYSUIS|eA ) puvoydijzaa pyouldy 
qsnsny Ajrea — Ayn pr (WIeYSsUIS|eM ) Duvaupjspo pyouldy 
Jaquiaydag piu — Ayn aye] (snavuurq) puplipuvjos pijouldy 
ysnsny avy — Ang pr (snovuur’y) pyjauosisy vyouldy 
Joquiaydag piu YOLulay] DUDIIPDA DPJASUH) 
Avy plu — Avy Ay.tea (WUeYSUIS|eAA ) DUOSALO DAaJUaXapNnas 
Ayng oe] []oMog pupunjs0fun DAIYADAAZ 
Jaquiaydag pri Z (Woy.way) puounjiof piaydv.i1az 
Ayn aie] URLUdd1 2 BAINNIN sisuappuvs v1aydvs1aZ 
oun ae] — oune piu YOLIULOH, pjoun[pp piuouosdéy 
A[ng oye] Z (SUdUa[D) pupjoo1ayps puouosdk&y 
Ajng aye] — oung pri YOU sluoynjysqns puouosdéH 

pollog 14314 So]ION Joyny saioads snuan Aprurey 


“Apnys 1edX-99.1Y) OY} QAO ape SUOTDAT[OI UO pase SI PoLiad WYSIPY “ISIP{IOYS (E€8H]) [B19 SOSPOH OY} SMOTI[OJ 


exe) JOYSIY UIYM IoUANbas satdadg “(GGG ) UISUIISIFY] JO IVY} SMOT[OJ UONLIYISsL]d JOYSIH “VUOG]Y ‘Oye poomyYoNoOL, Aeau payda][oo satoeds viaydopidayq jo si] (panunyuos) *Z ATAVL, 


(oe) 
ae) 


LEPIDOPTERA NEAR LAC LA BICHE 


POHL, LANGOR, LANDRY, and SPENCE 


2004 


Avy] Ay.rea (TDYTVM ) DIADSAN DIIKT 
Ayng prtu — sung pr (snovuur’y) DIADINJAG UOISI 
Isnsny Apia — oune ey] (aquany) DIADIJAIOd DIUUADOGOJOLd 
aun Apiea — Avy pr (ADT[NULIIFFIYIS wz stusq) piapnosndasd sidosjoq 
Ajng prt — Aeyy oye] (souany) DIADAAD] SISdOPIA] 
Av], prul (194[BMA ) DIXAJAIJU VDAN|DYJaV 
oung Apiva — Avy pr (TDATCAA) ppusoxa DIUOpyoyliCQ 
oun aye] (jayouodnq) piuvjsaddi DIUWPASIG 
Ayng pruu — sung Apiea (aouqnH) DIADUSIS DIADIDIN 
Ayng prtu — Aeyy pri ({jes1eog) DIDAIIS]N DIADIDW 
isnsny Apiea — oune aye] (1dYIVAA ) DIVIIDIIG VIADIDW 
ysnony Ayia — Ayne prw (UBLUSIOAq) DIADIIAO] DIADIDIN 
Ayng ayey — Apne pr (S1aquny,) DIVIUUNAG VIVID 
Ayn prtu — sung prut (as[ny) SIVUIGAIA AUDIO! — WUILUOUU — depLNOWIOIH 
Ajng prut — ounr 3e] (JOyTeAA) DASOA VIALCQ 
Ajng pri — Aevyy oye] (preyoed) pvauipig vuvdas1q 
ysnsny Apiva — Aevyy aye] JONTCAA vypno1v Duddaq 
oune Ayiwa — Avy] prul (aguany) suapnd paypaying 
Ang prut (a9uany) saplosoydoJDUkd DANDAYIJOPNAS J 
Ayng prtu — sung pr (asson) DIdIUIS AUKSOAGDH oeprurdaiqg 
ysnsny Ayiewo — oune aye] (Ainiq) SIUMAYIAD DIYIAD]ISDG 
snsny av] — oune ae] (qepoy) 1AIg] NU SIDISY 
aun aye] — 19q0190 Aye (snovuury) pdoyup sijpyduan 
Avy] prt — Aeyy Ay.ea (spreMpy) SNAKIDS DIUOSA]Og oepryeyduAnyy 
oune ae] ({BAnpsiog) DINJUAUD SaL1aagq ovpiuaroh'] 
aunf 31e] sLuey DaIVAsI]O SAI PL 
Aqng aye] — Avy oe] ueplor 2 P[LYOsyIOY sisuappuvo o1piddg ovprluorideg 
ounf oe] (sted) uowanppod snjpydados1ajADD oepruiodsay 
ysnsny Aiwa — A[ng pr (pyeusaj) ajpusisoquinjd Dwus1JSOjJsltoy) 
qsnony Aiea — oung ye] (AAV) SISA] DAP| 
Ajng Ajiva — Avyy oye] pieyorg Sypas1og DISNDAKG 
oune Ayia (9]01D) sypoiu visnvskg 
Ajng pruu — sung pru (jasvujny) DIDUOLOD DIUAVJIA] Yd 
oune ae] IO]]9Z, sypynoave vyspdsisad 
Aqne oe] (jaseujny) DIDpINd SISassaaq 
ounf prut (susua[D) sypnovu xudodvsvg 
ysnsny Ayiea — oun pr (TDyTeM ) sypsaqygo vyyouas 
Ajng aye] — Ayng pr (OJIeay) Dyjajoundissop visvipad 
polled Ws Joyny sotoads snuan Ayre] 


‘Apnys IeaK-daIY} IY) JOAO IPL SUOTIDAT[OI UO pase SI PoLtad IYSI]T4 “ISIP{IOYS (€86]) ‘Te 19 SOSPOH oy} SMOT[OF 
BXe] JOYSIY UIYIIM Jouanbas satdadg (GGG) UdSUDISITY JO JY] SMOT[OF UONVOYISse[O JOYSIF{ ‘eMOQLY ‘axe poomyonoy Ieau payda]joo sorseds esajdoprdayqT Jo ysvq (panuyjuod) J ATAVL, 


THE CANADIAN FIELD-NATURALIST Vol. 118 


538 


Aq ae] = Avy Oe] OMS” DIIDALJad DUBWIOLIPA FT 
isnsny Ayre qS[0H NB1093 DIAKWA] 
Ajng piu — ounr Apvo (AO][NULIDJJIYIS wz stusq) pvaappis piadoydyoy 
ysnony Ajiwo — Ayne pre (as]nH) DUIJAX S1YJ1NGY 
ysnsny ae] — Ayn pr (JOP) pypupjdxa siyyjng 
ysnsny Apia — Apne pr (AOTB AA) pivsndoad siyjyjny 
ysnsny Ayes — oune prt (s[ny) DSOWAOL DULOAJSSKG 
ysnony Apiva — Avy aye] (aguony) DIVDIJISAIY VUOAISSKG 
ysnsny oe] — ounr oyv] ({[esae9d) DIVIIYIDM DUOASSKG 
Ajng piu (jadvujny) DIDIUNA] DULOASSKG. 
Joquiajdag piu — ysnsny Ayia (snovuury) DIDANI DUOAISSKG — DVUTUDIB'T — aepLauoayH 
ysnony Apiwo — oung pruu (A9[YOSOP) piuppisidf pjndorsy 
Ayng ajyey — oung oye] (JOP) piapjount pindoog 
Ayn piu (as[ny) pypjjaoup vindoos 
qsnony Ayteo — Ayn pr (YIoMeH) pippunoquyy vjndoasy 
Ayn aye — Avyy oe] (sguony) piupulnpuad psoydojrKy 
oune aq] (paeyord) pypuUuadopunjod Dapp] —-—- IWULY.ID}S — 9epLNawUo0ayH 
ysnsny Apes — Apne aye] (AdJJBYOS-YSLIOH ) DIADISISAL DAWVIOIVUAN 
qsnony Ayre — Apne prt (SLI) DIADNIDUL DKIIS 
Jaquiaydag pit — jsnsny aye] (s9uaND) DIAD]JAISY DUIPQuUIVT 
Ayn prtu — Avyy aye] (a9uany) DIADAOAIMIND Yulsag 
Ayng ayey — oung pr TOYIRAA vIvsiAlp VjaduUvy 
oung aye] — Avyy ov] (aguany) DIADJOOID SIPOSY] 
sung oye] — Avy pri (souony) DIADSOSO]Yyd SIPOSV] 
ABA OR] (snovuurq) pidpsaapnd siposv]d 
aun aye] — oune Apvo JOYICAA DIVUNUAAap DUAaUDIaW 
ysnsny Apiwo — Avy aye] douany DIADUOIDUI DUAUDIaW 
oune Apwo — Avy pr IOYIRA piapaydig]p piuajas 
Joquiajdag pri — ysnsny Avo souany DIADUSDU SOWOUUT 
ysnsny oye] — Apne pr (souany) vypjsad papduvy 
Ayn Ajrwo — oung prtu (JO4[CM ) DIADISAUOY OLA 
ysnsny Ayes — Ang pri (Ain.iq) pjadsos adxjoyjuvx 
ysnsny Apiea — oune avy] (aauany) DIADUIAS 1] DUAD]YIN| 
Av IR] (our) DIADUISADU DUAD]YINA 
Ayn piu (touqnH) DIADSNIGO DUAVDJYINT 
Ayn piu — oung pr sauany DIADJOIIDA DABQDI 
ysnsny Ajiea — oune pr aouany DIADUAYJAAA DIAQDI 
19q0190 Aj.rea — Jaquiaydag prur (SLR H) DIADIPY SIUUDAT 
poliog ISIy SO]ON Joyny satoads snuan ATTUue,y 


“Apnys IdA-da.1Y} OY} IOAO pel SUOTIAT[OI UO pase ST polad IYSIPY “ISIPJIIY (€Q6]) ‘Te 39 SASPOH IY) SMOT[OJ 
BXL] JOYSIY UIIIM BoUONbas sarsadg ‘(GGG |) UASUAISLFY JO JY) SMOT[OF UONVOYTISsL]O JOYSIF{ “eOqLY ‘Oye poomyonoy, teau payoay[oo saroads vsaidopideyq jo asi] (panuyuor) ‘7 ATA], 


fag) 
va) 


POHL, LANGOR, LANDRY, and SPENCE: LEPIDOPTERA NEAR LAC LA BICHE 


2004 


Ayn Ayrea — Avyy prut (SLI H) DUDILIAWUD DUISapo]]|AYq oepiduresoise’'] 

isnsny Apes — oune pr pieyorg DIDAOUWD DIZZ1]]VIJ aeprUuRsip 
oune av] IOYICA pIdIIBIAIU DAOYdOGOT 
oune pri — Ary Apiea (1OMTVAA ) DIDANJYOAID DAVPYID 
ounr Ayiea — Avyy pr (1O¥TVAA ) DIADIUN] DADPVID 
Avy piu (preyord) DIDPIAIA SISDIV 
ounr Ayiea — Aevyy prul pieyorg DIVIVISOIOADA DIDAYNAN| 
oun ae] — sung pr JOYE piavoyup pigayndny 
ysnsny oye] — une pr (as[nH) DID Jas vioayndny 
Ayng prtu — ounr Ayiea (as[nH) pasnfsad pigaynldny 
ysnsny Apiva — Avyy oye] Aepolqnoqg ppjiuissy pigayydny 
Ajng ayey — Avypy 21] (touqnH) DIDAAIDS DIDAYIAN|A 
oun ae] — Aeyy oye] (YJIOMP}) pyposnfqns visayjidny 
Av P| (aehq) pipiquinjoo pivayndny 
ysnony Aiea — oung aye] (JD4TPAA) poipuau aydpqny 
19q0}90Q Ape (as[nH) pypaonag vsajydosad¢Q 
Jaquiajdag piu (Uasneyyog) pipuumnynd vylssdy 
Ajng prt — Aryl 312] (s9uaND) DIDIPAOG]D DIZAPOYIIAT, 
Ayng pri — Avyy pr (aeAq) 1jpsavad pisnua, 
ysnsny avy — Avy oye] sing DIIAQUIDI DISNUAA 
ouny 9}e] (1O4TRM) DAiafiqyy DYAaIpAH 
isnsny ae] — Ang pru OulsseZ pypuidjp sdajydouaz 
Aqng aye] — Avy pr (aguond)) vypipawaaqur pidydny 
Ajng oye] — ung pr (JOT [NJA) vpUsayp aoyssdy 
ysndny Ajiwa — Avyy pru (aguany) DIDAJSNIVD] OYLOYJUDX 
Ajng piu — Avy oye] (PID) DIDSNALA, POYAOYJUDX 
isnsny Ayiea — Ang piu (aadsq) DIADAO]OIAP BOYAOYJUDX 
isnsny Aiea — oung AT Ivo IO[ARL, DIADSSOL JOYAOYIUDX 
Ayng ae] (gouaNy) DIVNp1 aOYsOYJUDX 
ysnany Aiea — sung aye] (1dJJVYOS-YSLLIOH ) DIADSDAQD AOYAOYJUDX 
ARIA AL] (TOTEM) Dpsafyjnul vajayUy 
oune Ay.rea — Avy pr souony DIVIPISDA DAJIYUY 
Ajng aye — Aqne Ayaea (JOC ) DIVIDSDG YULOZ1A Aq 
Ajng prut — sung pru (AD][NULIAJJIYIS 2 studsq) DIpNjan] viuvs.ivds 
Ajng piu — Avy Apivo (suony) DIDIPLIYNA DINA|OSAW 
Jaqo1oQ Aira — Jaquiaydag piu (aguony) pypyisapy vsoydisy, 
ysnsny ae] — Apne aye] (Siaquny,) DIDIAN{ DUAULOLIPAH] 
Ajng ayey — Avyy oe] (194014) DIpAAGNA DUAUWIOLI PAL] 
Ajng aye] — Avyy oe] (ADT ) pIpIOUNUAL DUAWOLIPAL 

pollog SI SO}ON JouNy satoads snuan Ayre] 


‘ApNys IdK-9dAY} DY} JOAO IPLUT SUOTIITTOS UO pase st PoLtad IYSIpy “ISIP{IOYS (ERG) ‘Te 19 SOSPpOH oy) SMOT[OF 


BXe] JOYSsIY UTyIIM doUENbas satdadg (666) UASUIISITY JO IVY) SMOT[OJ UOTVOYISse]O OSI “ViOq[Yy ‘Oye poomyonoy, 1wau pajoa][09 sardeds vsgydopidsy jo jsv] (panuyuod) *7 AAV, 


Vol. 118 


THE CANADIAN FIELD-NATURALIST 


540 


Ayn oyey — Apne pra aauany sypnbuidosd pinay OVUI[NALY — oepINIoON 
ysn3ny Apiewo — Apne pro (UIUS “g'f) pauliyovisf sapouadiy seurueutsdans — aepmnjoony 
Ajng piu — sung oye] (aauqnH) SYDINBUD SIYI]Vq 
Ajng prut dauany SIPULIpvADI Duldalg 
Ajng a1] vd (UIs “g'f) NUDYUDY VjOISOUay|PUd 
ysnsny Aiea — Avy] oe] 01D sypasjad vINOIKYD 
ysnsny Ayivo — Apne pr (yt “g‘f) pq]vIN] VYyIwUsO]IUDZ 
Ayn prt — Avy oye] (IDTV) sypsnupskd punydouanjpyd 
ysn3ny Ajiwo — A[ng prut ¢€ Jouqny pinwav eau saisads Mau VIP] 
ysnsny Ayia — Ayn pr Jouqny pinuav VIP] 
ysnsny Apieo — suns oye] (aguany) SIJDIIAAWUD DIP] BUNULULIOF — OepInjoOoN 
ysnony avy — Avy pr (TDAP) puppisidf DjoajotN — seuldt1yjo.LIeg — aepinjooN 
Ayn ayy — Apne pra (TDYIPMA ) pypispyd palyIksvg 
Ajng oyey — Apne pri (ysnouunqgdsyy 2p soureg) SUDBDA DAIYIASDG depILUeULAT 
oun aye] — oun piu SLLIeH ppjnoput vduma20ydoT 
Ayng piu — oune pru (SLLIeH) souayjipd DYIAVID] 
oune a}e] — Avy ae] JORMA SUD]IUNSSD DIGOJDUSDAY 
qsnsny Ayiea — Ayn piu (O[[IAQUSAJ-ULIOND ) 1ajuoza] vojdpy 
ysnsny Apivo — Ang prur pieyord DID|G]Y vIsuauia]D 
ysn3ny Ayres — Ang pr (2}01D) 4OJONIG DUA] oeplyno1y 
ysnsny Ajiewa — une aye] (01) saplouyda] vanziyos 
Ajng Ay.ea (yyUIS “A‘f) STUAODIUN DANZIYIS 
oune ae] (uospny) DISAPOU DINIANA 
oune ae] — Aeyy ae] (eAnpsiog) pUulApuadojoos VINIANA 
oun aye] — Avy oye] (touquUr'T) SIDJUAPIIIO VINIAN 
Avy Ayivo (01D) laujuy pisiydnjpH 
Avy, RY] — Avyy pre uospny pynopuan visiydn]H 
ysnsny Ajava — Aeyy ey] IONE stuoluajdas visiydn]y 
oun Ajrea — Avyy oye] jovin pip) dus DJUOPOJON 
Aqne piu — Avy prur pieyorg DSOUL DISOAY 
oune pruu (YJIWIS “A'f) DSOgq1s DIDPPN 
oung aye] — Ary oie] (IOyTRM) sypo1dv p1ajsoj]D 
sung aye] — Avy oye] (01D) DSOBIAJS DAAJSO]D 
Ajng avy — Avy ayy you DUSISOG]D DAAISO]D oepnuUopo}ON 
oun ae] (Sinquia}0y) mp8 Saja 
isnsny Aiwa — Avy pr AQAry IAS1499 SNYJULLAWUS oeprsuryds 
ysnsny Apiewo — Apne pra JouqnH DIAISSIP DULOSOIDIVDIN 
pollag IWSIpy SION Joyny satoads snuan ATTUre.y 


‘ApS IRIA-J9IY} IY) IOAO OPBU SUOTIAT[OS UO paseq st PoLtad IYSIPY “ISTPIIYO (€QG]) “[e 9 SOSPOH BY) SMOTIOY 


Bxe] JOYSTY UTYTM dUINbes sa1sadg *(6GGH]) UBSUDISITY, JO JY SMOT[OJ UOTBOYTISsKIO JOYSIF “VUOQTY ‘oye poomyonoy, teu payoayfoo saroads viajydopidayq jo isv] (panuiuod) ‘7 ATAV], 


54] 


IRA NEAR LAC LA BICHE 


NCE: LEPIDOPTI 


4 


POHL, LANGOR, LANDRY, and SPE 


2004 


ISNSNY av] 


(aakads) 


pupoiiauy paodiyduy 


Isnsny ayey — Ayn pr (aouqn}) DIIAUSNS SUYINSDADE 
Jaquiajidag prw — jsnsny ar] (JOYIVAA) VIDIO}! DISCO 
Jaquiaydag piu — ysnsny aye] (sauany) DIVIIDU DISICQ 
snsny Ayiea — Ayng ae] (ujTwIS) pID]IBOI Daubdy 
Ayng aqey — Avyy Ayiea (JOMTeAA) ppowuos vaupdy avuluapeH — oepinjooN 
jsnsny ae] — Apne ey] (JOMPA\) pxiful Duapoyouoryy 
snsny ayy — Apne pra (9}019) DSLAJSIPDG DUaPYYOUlo yy OVUIT][NOND — oepInjoOoN 
ounr piu (TOYTR A) DIDUSISIA] DUUAULISIAAD FY] 
aune piu — Aryl aye] JORMA DSSatdull DIJUOLIY 
oun ae] — sung Area JOYE Djajdul vjgU0LIy 
snsny Aiea — APY] Ae] (aguany) SIS DAL DIDIUOLIY 
Ajng piu — Aevyy aye] JOYE VASUs DIMUOLIY 
snsny Ajiea — oun pr aguony DIDIJOUUL DIIIUOLIY 
aune ae] — dune Aye douony DUldna DIIUOLIY 
Ayng aqyey — Aryl ore] OID Jawdf viydvy —- aeua1uosdVy — ovpIndd0N 
Ajng aie] — oun prur (a99uaND) DINpIG]Y VIPOIVYINT dvUIO.NSNY — sepinjooN 
ysn3ny Apia — une ae] douonh nupujnd pisn}d 
Ajng pr (AqIry) vinsuvpjoad DyddAsUuKsS, 
Aqng aye] — Apne pr (Insusjongd) spyp pydvssuasy 
ysnsny avy — Apne aye] (Q]01D) DUB ISIPIA1A DYADABUKS 
A[ng ae] (9019) pidiagsoja0 pydvssuasy 
Ayn ayey — oung aie] (1DYTRAA) pidup pydvisoiny 
Ajng prt — oune oye] (UOSUTGOY 2 901) pddpu vydpisojny 
ysnony ary — Ang aye] (suoydays) pypnovuig vydpssoiny 
oun aye] — oun pru Insud;OnNO ppigqna pydpasojny 
Ayn ae] (sound) saproskyoay) XKsajdosoydsogq 
Ayn ae] (Q101D) psowsof pyduicuvsKy) 
ysnsny Ajiea (anyioqQ) DP]alauUsa DISKAYIA] Og 
Ajng ae] (9101D) saploasav DISKAYIVDIG. 
Ayn prt — oun 39e] aouonty S1JUAIN DIOISOAGY avuUlIsN[d — sepInjooN 
snsny ae] — snsny Ajies 01D DIIYAAMUIS DIVIOIDD 
Jaquiajdag piu — jsnsny Ayiea spiempy SIASIAG DIDIOIDD 
Jaqo1oQ Apiea — Apne aye] JOC pentiun pjps0jvyD 
Jaquiajdag prut —ysnsny Ayieo JOC DJIAL DIDIOIDD 
ARYA] IE] (WJIOMeH) DINISNISSDAI DULIBANUID IVUT[RIOJVD — OepINIION 
Avy pruu STeY ynuny Duaday 
isnsny aye] — Ang pra (Aye ) sypjoipa puaddy 
Ajng prur quis piupwojyp puadA yy avuruadAY — oepinjooN 
polled ISt14 loyny soroads snuan Ayre 


‘ApNys IdK-IdAY} OY} JQAO IPRUT SUOTIDOT[OD UO pase SI PoLtod IYSIL4 “ISIPYOOYS (€Q6]) [P19 SASPOH IY} SMOTIOJ 


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Vol. 118 


THE CANADIAN FIELD-NATURALIST 


542 


Ayng oye] — Apne prow 

ysnsny Ajiva — Apne pr 
ysnsny yey — Ayn pri 

oun Aj.ro 

oun Ay.eo — Avy plu 

oune Ayrea — Avyy Apieo 
Avy pr 

oun Ayre — Avy pra 

oune ae] 

Ayne ayey — Apne prot 

ysn3ny Ajiea — sung aye] 
Ayng pra 

Ajng ae] — oune aye] 

ABI, OP] 

oung pri 

Ayn oyey — Apne pra 

ysnsny Ayjieo — oune piu 
snsny oe] — Isnsny Aj.ea 
ysnsny aye] — Isnsny Ayia 
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Jaquiaydag piu 

snsny ae] 

Jaquiajdag piu 

Jaquiajdag prt — ysnsny Aya 
oung Aye 

oung Ape 

oung Ajiva — Jaquiajdag piu 
19q0}9Q Aj.iea 

Jaquiaydag pr — \sndny ae] 
Avy] plu — Joquiajdag pri 
oun ayey — Avy oe] 

Ayn pr — oung prut 
snsny Ie] 

Ajng piu — Avyy oye] 

ysnsny oe] — snsny Ay.ea 
Jaquiaydag piu — A[ng prur 
Joquiajdag piu — Ajng oye] 
isnsny Ajaiva — Ayn pr 
oune aye] — oun Avo 


polled 1Yyst[4 


SOON 


(91019) 

(yMUIS) 

(Q]01D) 

(aguany) 

(21019) 

(aguany) 

(YIU ) 

(UOSTIIOJ]) 

sguony 

(Q]01D) 

(s9uony) 
(suaydays) 
(uruuefuag 2 sou) 
(1O4TBA\ ) 

(aguand) 

(e9uan5) 

(s9uony) 

(194[2M\) 

(01D) 

urueluog 2 souieg 
(suoydays) 
Qpeis.i0197) 
PIOPTNA 2 surejuoje’T 
(UOSUIGOY 2 9101) 
01D 

(y}IUIS) 

(UOSLLIO[\) 
UOSLLIO[ 
(UOSLLIOJ[) 
(UOSTLIOJ]) 
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(TOTEM ) 

(soueny)) 

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543 


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Aqng ayey — Ayu prur 
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isnsny aye] — Apne aye] 
snsny Aiea — Avy Aprea 
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Aqng ayey — Avy 1e] 
Avy prul 

ysnsny Aiea 

A[ne ae] 

snsny Aiea 

Apne are] 

isnsny aye] — Apne aye] 
Ayn are] 

Ajng pr 

ysnsny Ajiva — Ayng prur 
isnsny aye] — Apne prw 
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pouled 1yst[4 


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(21019) 
(UOSTIIOJ]) 
(21019) 
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(sguony) 
(TONTEAA) 
(9}01D) 
(UOSTLIOJ{) 
(touqnH) 
(TOTEM) 
(uaqjaus) 
(M019) 
(snovuurT) 
(UOSLLIOJ]) 
(snovuul’y) 


(sniotiqe,y) 


JoyINYy 


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puaypsad Sols DJa,any 
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pylsadul viqouyovd 
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DUDIUDUULAOU DYSIX 
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544 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Gelechia lynceella 


FicureE 2. Microlepidoptera collected near Touchwood Lake, Alberta. All are new Alberta records. (ruler = lcm). 


The study site lies within the ranges of 67 butterfly 
species (Bird et al. 1995); many of these certainly occur 
in the mixedwood habitat but were not collected there. 
Some nocturnal moths, including some species of 
Gelechiidae and Oecophoridae (Hodges 1974; Miller 
2000), are not attracted to UV light so they would be 
undersampled as well. If other moth species occur in 
the area but were not collected, they are probably quite 
rare, or may be extremely localized in particular micro- 
habitats, which were not adequately sampled in this 
study. 

Macro-moths comprised a greater proportion of 
individuals than of species, whereas the micro-moths 
were particularly species-rich, comprising a greater 
proportion of species than of individuals. These pro- 
portions (Table 1) were similar to those previously 
reported for the province of Alberta. In his list of 
Lepidoptera of Alberta, Bowman (1951) reported 40.8% 
micro-moths (779 species), 50.9% macro-moths (973 
species), and 8.3% butterflies (159 species). In the most 
recent list of Lepidoptera of North America, Poole 
(1996) listed 49.3% micro-moths (5743 species), 43.9% 
macro-moths (5114 species), and 6.9% butterflies (801 
species). The proportionally higher representation of 
micro-moth species in the current study compared to 


Bowman’s 1951 list reflects the recent increase in 
knowledge of the group, including the description of 
many new species. 

A total of 35 positively identified species are new 
records for the province, and one (Acanthopteroctetes 
bimaculata) is a new record for Canada (Table 2). 
Micro-moths (Figure 2) make up 34 of these new 
records; a further five micro-moth species and one 
macro-moth species represent undescribed species. 
Details for some of these new records and new species 
appear below. 


Acanthopteroctetes bimaculata Davis (Acanthopteroctetidae): 

This primitive moth is known previously from California 
and Oregon (Davis 1978). The current record represents the 
first report of this moth, and of the family Acanthopte- 
roctetidae, occurring in Canada. Nothing is known of its 
biology. 


Caloptilia anthobaphes (Meyrick) (Gracillariidae): 
Previously reported from northern Ontario (Forbes 1923). 
Nothing is known of its biology. 


Caloptilia betulivora McDunnough (Gracillariidae) (Figure 2): 

Previously reported first only from Nova Scotia (McDun- 
nough 1946) and since been reported from Quebec (Hand- 
field et al. 1997). It likely occurs across the boreal zone in 
Canada. Larvae feed in the folded leaves of birch. 


2004 


POHL, LANGOR, LANDRY, and SPENCE: LEPIDOPTERA NEAR LAC LA BICHE 


Chytonix palliatricula 


FiGurE 3. An assortment of macrolepidoptera collected from mature aspen forest near Touchwood Lake, Alberta (ruler = 1 cm). 


Caloptilia canadensisella (McDunnough) (Gracillariidae) 
(Figure 2): 

Originally described from Nova Scotia (McDunnough 
1956). It has since been reported in Quebec (Handfield et al. 
1997; Landry and Landry 1992), and is probably widely dis- 
tributed but uncollected across the boreal forest. The larvae 
make large blotch mines on the leaves of Bunchberry (Cor- 
nus canadensis). 


Caloptilia coroniella (Clemens) (Gracillariidae): 
Previously known only from the midwestern U.S.A. 
(Forbes 1923). Larvae feed on birch. 


Caloptilia stigmatella (Fabricius) (Gracillariidae): 

Not reported from western Canada, but GRP [Greg R. 
Pohl] has collected it quite commonly in Alberta and Saska- 
tchewan. Larvae feed on willow (Forbes 1923). 


Argyresthia abies Freeman (Yponomeutidae): 
Reported in eastern Canada only as far west as northern 
Ontario (Freeman 1972) but is probably widely distributed 


across western North America in the boreal forest. It is a 
twig borer on Balsam Fir (Abies balsamea). 


Coleophora corylifoliella Clemens (Coleophoridae): 
Previously reported only in eastern North America (Forbes 

1923). Larvae there feed on Corylus americana; in the west 

they probably feed on Beaked Hazelnut (Corylus cornuta). 


Coleophora duplicis Braun (Coleophoridae): 

Previously known from eastern Canada and the midwestern 
United States (Forbes 1923: Handfield et al. 1997). It feeds 
on the seeds of Aster (Aster spp.) and Goldenrod (Solidago 


spp.). 


Limnaecia phragmitella Stainton (Cosmopterigidae) (Fig- 
ure 2): 

A holarctic species. The closest it has been reported to 
Alberta is Wyoming (Hodges 1978). However, collection 
records (CNC, NFRC) indicate that the species is transamer- 
ican. Recent collections made by GRP indicate that it is com- 
mon in western Canada, at sites where its host plant, Com- 


546 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


cronicta innotata 


Ficure 4. An assortment of macrolepidoptera collected from old growth aspen forest near Touchwood Lake, Alberta (ruler 


—yluemi)® 


mon Caitail (Typha latifolia), is available. Larvae feed on 
the flowers and seeds. 


Coleotechnites blastivora (McLeod) (Gelechiidae): 
Reported only from the type locality of Gaspe, Quebec 
(McLeod 1962). It probably occurs across the boreal zone. 


2000 


Noctuidae 

HB Geometridae 
other macro-moths 

{) micro-moths 


no. specimens 
_ 
oa 
o 
o 


June July Aug. Sept. 


trapping date 


FiGure 5. Abundance of major Lepidoptera groups collect- 
ed in UV traps through the 1994 trapping season. 


Larvae are needle webbers of White Spruce, occasionally 
mining within the needles. 


Gnorimoschema septentrionella Fyles (Gelechiidae): 

Miller (2000) reports this species only in eastern North 
America, as far west as Minnesota. It is a stem-gall maker 
on asters (Aster spp.). 


160 


©) Noctuidae 

GN Geometridae 

__| other macro-moths 
{) micro-moths 


120 


no. species 
co 
oO 


sy 
Oo 


June July 
trapping date 


Aug. Sept. 


FiGure 6. Species richness of major Lepidoptera groups 
collected in UV traps through the 1994 trapping 
season. 


| 


2004 


Aethes promptana (Robinson) (Tortricidae — Tortricinae): 

The Touchwood Lake record represents the first record of 
this species from western Canada. It was previously known 
from eastern North America as far west as Wisconsin (Sabourin 
et al. 2002). A prevoius record from Washington (Razowski 
1997) was based on a misidentification. Nothing is known of 
it’s feeding habits. 


Wockia asperipunctella (Bruand) (Urodidae): 

As noted by Landry (1998), this Touchwood Lake record 
represents the first report of this species, and the family Uro- 
didae, in Alberta. This family is a recently recognised addition 
to the North American fauna (Heppner 1997). Larvae feed on 
Trembling Aspen; adults tend to fly during the day or early 
evening. 


Alucita lalannei Landry & Landry (Alucitidae): 

Until recently, this species had been combined with two 
other species in the genus, and referred to collectively as Alu- 
cita hexadactyla (Linnaeus) (Bowman 1951; Hodges et al. 
1983; Poole 1996). However, the true A. hexadactyla 1s restrict- 
ed to the Old World. In their description and treatment of A. 
lalannei, Landry and Landry (2004) designate a specimen 
from the current study as a paratype. This species is known 
from Ontario, Manitoba, and Alberta. Adults overwinter, 
and can be collected from May to September. The larval 
host plant is unknown, but Landry and Landry (2004) report 
Lonicera spp. and Symphoricarpos spp. as likely candidates. 


Parapoynx maculalis (Clemens) (Crambidae): 

Previously known from eastern North America, only as 
far west as Lake of the Woods, Ontario (Munroe 1972). How- 
ever, recent collecting by GRP indicates that it is widespread 
but extremely localised in Alberta and Saskatchewan. Its larvae 
are aquatic, and are reported by Munroe (1972) to feed on 
several species of water lilies (Nuphar, Nymphaea, Brasenia 
spp.). It likely feeds on other plants as well, since it has been 
collected recently in southern Alberta, outside the distribution 
of the aforementioned plants (C. D. Bird, personal commu- 
nication). 


Idia new species near aemula Hiibner (Noctuidae — Her- 
miniinae): 

Usually mixed in with /dia aemula Hibner in collections. 
It was reported in eastern North America by Rings et al. 
(1992), and misidentified as Epizeuxis concisa Walker by 
Forbes (1954). It remains undescribed and has not been 
reported previously from Alberta, although it is common 
across the boreal region. It has been reported to feed on the 
needles of a variety of conifers (Rings et al. 1992). 


Phalaenostola hanhami (J. B. Smith) (Noctuidae — Her- 
miniinae): 

Reported from eastern Saskatchewan as far west as Regina 
(Hooper 1988). Nothing is known of its feeding habits. 


The only similar published study of boreal forest 
Lepidoptera that the authors are aware of is that of 
Morneau (2002), which sampled primarily macro-moths 
near Peace River in northwestern Alberta. In that 
study, 293 species were collected over three seasons, 
including 278 species of macro-moths. Although most 
macro-moth species were common to both studies, 
43 species were unique to the present study, and 115 
species were unique to the Morneau study. The pres- 
ence of so many unique species suggests some habitat 


POHL, LANGOR, LANDRY, and SPENCE: LEPIDOPTERA NEAR LAC LA BICHE 


547 


specialization within the boreal mixedwood region. 
Several of the species unique to the Morneau study, 
most notably some Arctiidae species, represent an 
incursion of cordilleran species into that area. 

Several Lepidoptera inventories of provincial parks 
in the boreal mixedwood forest of northeastern Alberta 
have been carried out (Schmidt and Pohl 2001*; 
Macaulay and Pohl 2002*, 2003*). Although these 
studies collected from 138 to 295 species, they were 
each based on brief sampling periods, so they missed 
significant proportions of the fauna. Catches from the 
current study shared approximately 50 to 60% of the 
species collected in those studies. 

In general, the moth fauna of the boreal forest appears 
to be less diverse than that found in other forests. 
Summerville and Crist (2002) collected 512 species 
of moths in deciduous forests in Ohio. Thomas et al. 
(1998) collected 624 species of macro-moths in Fundy 
National Park, New Brunswick, which lies in the Aca- 
dian Forest Region, as described by Rowe (1972). 
Both of these forest types are more diverse botanical- 
ly than the boreal forest, and have more moderate cli- 
matic conditions (Rowe 1972). 

Inventories of this nature are a necessary prerequi- 
site to understand impacts of forestry practices and 
climate change on biodiversity. However, there is a 
paucity of similar studies, largely due to the lack of 
required taxonomic expertise. This argues strongly for 
increased support of systematics research in Canada, as 
biodiversity issues continue to increase in importance. 


Acknowledgments 

Héctor Carcamo, Tom Clark, James Hammond, Rob 
Lucas, Phillippa Rodriguez, Peter Shipley, the late Tim 
Spanton, Kevin Sytsma, and Daryl Williams assisted 
with field collecting. The Alberta Land and Forest 
Service (Lac La Biche Region) provided field acco- 
modation, and the Alberta Research Council provided 
maps and assisted with site selection. Gary Anweiler, 
Klause Bolte, P.-T. Dang, Ron Hodges, Don Lafontaine, 
Doug Macaulay, and Jim Troubridge assisted with 
identifications. This work was funded by the Canada- 
Alberta Partnership Agreement in Forestry and by the 
National Science and Engineering Research Council 
of Canada (operating grant to JRS). 


Documents Cited (marked * in text) 

Macaulay, A. D., and G. R. Pohl. 2002. Survey of Lepi- 
doptera in the Canadian Shield Ecoregion of northeastern 
Alberta. II. 2001 Survey of La Butte Creek and 
Fidler/Greywillow Wildland Provincial Parks. Report to 
the Alberta Natural Resources Service and Alberta Nat- 
ural Heritage Information Centre, Parks and Protected Areas 
Division, Alberta Community Development. 61 pages. 

Macaulay, A. D., and G. R. Pohl. 2003. Survey of Lepi- 
doptera in the Canadian Shield Ecoregion of northeastern 
Alberta. III. 2002 Survey of Colin-Cornwall Lakes Wild- 
land Park. Report to the Alberta Natural Resources Ser- 
vice and Alberta Natural Heritage Information Centre, 


548 


Parks and Protected Areas Division, Alberta Community 
Development. 44 pages. 

Schmidt, B. C., and G. R. Pohl. 2001. Survey of the butter- 
flies and moths (Lepidoptera) of the Canadian Shield 
natural region of Alberta. Report to the Alberta Natural 
Resources Service and Alberta Natural Heritage Infor- 
mation Centre, Parks and Protected Areas Division, 
Alberta Community Development. 30 pages. 


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Bowman, K. 1951. An Annotated list of the Lepidoptera of 
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Chao, A., and S. M. Lee. 1992. Estimating the number of 
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Colwell, R. K., and J. A. Coddington. 1994. Estimating 
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Received 20 October 2003 
Accepted 27 September 2004 


Pilose Braya, Braya pilosa Hooker (Cruciferae; Brassicaceae), an 
Enigmatic Endemic of Arctic Canada 


JAMES G. HARRIS 


Biology Department, Utah Valley State College, 800 West University Parkway, Orem, Utah 84058 USA; e-mail: harrisji@ 
uvsc.edu 


Harris, James G. 2004. Pilose Braya, Braya pilosa Hooker (Cruciferae; Brassicaceae), an enigmatic endemic of Arctic Canada. 
Canadian Field-Naturalist 118(4): 550-557. 


Braya pilosa Hooker, Pilose Braya, has been poorly understood among North American botanists due to a paucity of fruiting 
specimens for study. This has resulted in confusion about the taxonomic position of the taxon within Braya, and has led to 
speculation about its generic status. An examination of fruiting specimens from the Royal Botanic Garden Herbarium at 
Kew, England reveals that B. pilosa is correctly placed in the genus Braya, and that it is a distinctive member of the genus 
deserving recognition at the specific level. I discuss the historical evidence that B. pilosa may not have been collected since 
1850 due to its extremely restricted distribution on the Cape Bathurst Peninsula of the Northwest Territories of Canada. I also 
present evidence suggesting that B. pilosa is diploid and may be a parent species to some of the more widespread members 
of the genus, all of which are polyploid. Its closest living relative is probably B. thorild-wulffii. 


Key Words: Braya pilosa, Pilose Braya, discovery, rediscovery, distribution, collections, rare species, Cape Bathurst, North- 


west Territories, Canada. 


Braya pilosa Hooker (Figure 1) is an extremely rare 
endemic of the Northwest Territories of Canada. It is 
perhaps the most distinctive and, at the same time, least 
understood North American member of a taxonomi- 
cally difficult genus. While some botanists (Harris 
1985; Argus and Pryer 1990; Rollins 1993; McJannet 
et al. 1995) have treated B. pilosa as it was envisioned 
and described by Hooker (1830), most have placed it 
into synonymy under B. glabella (Polunin 1959; Welsh 
1974; Porsild and Cody 1980; Cody 2000; Warwick 
et al. 2000*) or B. purpurascens (Scoggan 1978). Oth- 
ers have reduced it to infraspecific status within B. 
purpurascens (Schultz 1924; Hultén 1970), applied 
the name broadly to elements of B. glabella (Porsild 
1943, Hultén 1968), or ignored it entirely (Cody 1979; 
Mulligan 2002). To the best of my knowledge, there 
are no authentic fruiting specimens of B. pilosa in any 
North American herbarium. It is this lack of available 
fruiting material for study that is primarily responsible 
for the taxonomic confusion that has surrounded this 
taxon over the past 173 years. 

When he described the species, Hooker (1830) had 
only a single collection of flowering individuals with 
immature silicles from which to work and he was ap- 
parently uncertain whether or not the plant was actu- 
ally a Braya. He placed a question mark behind the 
genus name, lamented the lack of mature fruits that 
would allow unequivocal generic placement, and stat- 
ed, “...in all probability, it ought to constitute a new 
genus.” Without access to mature fruits for examination, 
Hooker’s suggestion of a new genus would indeed 
seem justified. The flowers are larger than those in 
any other Braya species (the petals in some tetraploid 
B. humilis are as long but not as broad); the immature 


silicles are ovoid-ellipsoid, unlike those in any other 
Braya with the exception of B. thorild-wulffii (which 
had not been discovered in 1830); and the styles are 
exceptionally long for a Braya, measuring over half the 
length of the ovary. In addition, Sir John Richardson 
(1830), who collected the type specimen in 1826 during 
the second Franklin expedition in search of a north- 
west passage, described the flowers as fragrant, with 
a smell similar to lilac blossoms. Fragrance has not 
been associated with any other member of the genus. 

Murray (1983*) and Scott et al. (2000*) suggested 
that B. pilosa is a misnomer applied in error by Hook- 
er (1830) to a Draba, likely D. corymbosa. However, 
Hooker’s original description and accompanying illus- 
tration of the taxon indicate that he is describing some- 
thing quite different from D. corymbosa. Hooker 
describes the valves of the immature silicles as “valde 
convexis,” and again later as “remarkably convex,” 
descriptions that do not accurately describe the dis- 
tinctly flattened valves of D. corymbosa. 

Fortunately, any doubt about the identity of Hook- 
er’s B. pilosa as a legitimate Braya rather than as a 
misidentified D. corymbosa 1s resolved by an exami- 
nation of authentic fruiting specimens. There is a herb- 
arium sheet at the Royal Botanic Gardens Herbarium 
(K) in Kew, England (Figure 2) that apparently in- 
cludes specimens from three separate collections (Sea 
Coast, Arctic, Richardson s.n., 1848; arctic coast, W. 
of C. Bathurst, Captn. Pullen s.n., Aug. 1850; and what 
are likely part of Richardson’s type collection of 1826). 
The herbarium sheet is so congested that it is impos- 
sible to be certain which individuals are part of the 
various collections represented, but all of them are defi- 
nitely members of the same taxon. Mature silicles on 


550 


2004 


HARRIS: PILOSE BRAYA: AN ENDEMIC OF ARCTIC CANADA 


Ficure 1: Illustration of Braya pilosa. 


55] 


Bp 


these specimens, even though pressed, are clearly 
ovoid-ellipsoid and perfectly match Hooker’s illus- 
trations of immature fruits of B. pilosa. In addition to 
possessing fruits that are strikingly different from those 
of D. corymbosa, the Kew B. pilosa specimens differ 
significantly from D. corymbosa in several other attrib- 
utes. The leaves are much longer, narrower, thicker, 
more obtuse, and less densely pubescent than those in 
D. corymbosa; some individuals bear a single cauline 
leaf while D. corymbosa lacks cauline leaves; the styles 
are much longer than those of D. corymbosa; and most 
importantly, the epidermal cells of the silicle septum 
are typical Braya epidermal cells, characteristically 
thickened and transversely or obliquely elongated 
(Harris 1985, Figure 2), in stark contrast to the thin- 
walled, irregularly shaped, and essentially isodiametric 
septum epidermal cells typical of Draba. Braya pilosa 
is unequivocally not a misidentified Draba. 

I have not seen any authentic specimens of B. pilosa 
that have been collected since 1850. It is possible that 
it is extinct, as Argus and Pryer (1990) and McJannet 
et al. (1995) have suggested, but I think it more likely 
that its distribution is so limited that it has simply been 
overlooked by botanists. Hooker’s (1830) description 
of the type locality of B. pilosa, “Mouth of Macken- 
zie River, lat. 70°” may be somewhat misleading to 
those who have looked for B. pilosa over the last cen- 
tury or so. Porsild, for example, made several collec- 
tions of Braya between the mouth of the Mackenzie 
River and Nicholson Island in Liverpool Bay, where 
Hooker’s location description suggests that B. pilosa 
should be found. Although Porsild initially identified 
his collections as B. pilosa (Porsild 1943), none of them 
show the distinctive ovoid-ellipsoid silicles, exception- 
ally long styles, and large flowers of this taxon and 
fall instead within the normal range of diversity in B. 
glabella. 

There is evidence that Hooker’s description of the 
type locality may be his interpretation of Richardson’s 
broad conception of the Mackenzie Delta rather than a 
precise description of Richardson’s collection location. 
In his account of the 1826 journey from the Mackenzie 
River to Cape Bathurst, Richardson (1828) repeatedly 
commented on the alluvial deposits and sandbars his 
party encountered in the area, and correctly attributed 
them to outwash from the Mackenzie. He clearly 
viewed the entire region of the Tuktoyaktuk Peninsula, 
Eskimo Lakes, and Liverpool Bay as a product of a 
historical Mackenzie Delta. It is likely, however, that 
Richardson was actually very near Cape Bathurst rather 
than the mouth of the Mackenzie when he collected 
the type specimens of B. pilosa. In fact, there is some 
evidence that all three known collections of B. pilosa 
(Richardson in 1826 and 1848; Pullen in 1850) may 
have come from the same stretch of seashore near Cape 
Bathurst, roughly 200 km northeast of the mouth of 
the Mackenzie River. 

John Richardson’s original collection of B. pilosa 
almost certainly occurred on 18 July 1826, as the east- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


ern detachment of Franklin’s second expedition in 
search of a northwest passage was approaching Cape 
Bathurst. This eastern detachment, under Richard- 
son’s command, was assigned the task of exploring the 
arctic sea coast by boat from the Mackenzie River 
east to the Coppermine River, then traveling overland 
on foot from the Coppermine River to Great Bear Lake. 
On 18 July, Richardson (1828) records stopping for 
breakfast “about eight miles” from the narrow passage 
between Cape Bathurst and the Baillie Islands. He 
writes, “The air was perfumed by numerous tufts of a 
beautiful phlox, and of a still handsomer and very fra- 
grant cruciform flower, of a genus hitherto unde- 
scribed.” There is little doubt that the “cruciform” 
flower Richardson is describing is B. pilosa. First, 18 
July is a date that an arctic Braya would be expected 
to be in a flowering rather than a fruiting condition. In 
addition, the dense clusters of large flowers on his type 
specimens would certainly be “handsome,” and Rich- 
ardson’s description of the plant’s fragrance and uncer- 
tain generic status matches perfectly with Hooker’s 
(1830). No other cruciferous species included in Hook- 
er’s Flora Boreali-Americana, which treats all of Rich- 
ardson’s collections from this expedition, fits Richard- 
son’s 18 July plant description as well as B. pilosa. 

Richardson collected B. pilosa a second time in 
1848, this time when the plant was in fruit. Captain Sir 
John Franklin, along with his ships, the Erebus and 
Terror, and their crews disappeared in the Canadian 
Arctic in 1845. Franklin’s ships carried sufficient pro- 
visions to last until the summer of 1848, but by 1847, 
when nothing had been heard from Franklin, the British 
Admiralty began mounting search expeditions. In 1848, 
John Rae and John Richardson were placed in com- 
mand of one of these search parties. Richardson’s 
orders were to search the arctic coast and islands east 
of the Mackenzie River to the Coppermine River and 
then travel overland from the Coppermine to Great Bear 
Lake, essentially retracing his 1826 journey. Unfortu- 
nately, Richardson did not mention his 1848 collection 
of B. pilosa in his published account. However, I think 
it is likely that this second collection was made at ex- 
actly the same place as the first. On 10 August, Rich- 
ardson was approaching Cape Bathurst again, twenty- 
two years after his first visit there (Richardson 1851). 
This is exactly the right season to find Braya in fruit. 
Given Hooker’s uncertainty about the generic status 
of B. pilosa due to a lack of fruiting material for exam- 
ination, it is inconceivable that Richardson would not 
have taken the opportunity to revisit his original col- 
lection location to collect fruiting plants which would 
allow a definitive identification. 

Braya pilosa was collected for a third and apparently 
last time in August of 1850 from the “arctic coast, W. 
of C. Bathurst” by Captain William J. S. Pullen. In 
1850, Pullen, also in command of an expedition in 
search of Franklin, attempted to travel from Fort Simp- 
son on the Mackenzie River to Cape Bathurst and then 
on to Banks Island. In mid-August he was in the vicin- 


2004 HARRIS: PILOSE BRAYA: AN ENDEMIC OF ARCTIC CANADA 553 


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FiGuRE 2: Photograph of mixed collections of Braya pilosa on a herbarium sheet at the Royal Botanic Gardens Herbarium 
(K) in Kew, England. Insert is an enlarged view of mature silicles. 


554 THE CANADIAN FIELD-NATURALIST Vol. 118 
MSW 129°00W 28°45 WBS SC IBMSW «IBOOW «= 1245W O12S0W SC 121BW12P00W Ss 128°45W «126° 
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FiGuRE 3: Map of Cape Bathurst Peninsula, Northwest Territories, Canada. The dashed box indicates the probable type 


locality of Braya pilosa. 


ity of Cape Bathurst, but found dramatically different 
ice conditions there than Richardson had two years 
previously. The pack ice was still so thick that he was 
unable to proceed past Cape Bathurst, and was forced 
to retreat back to Fort Simpson. On 8 August, Pullen 
(1979) recorded that his party “landed in a small bay 
about 7 miles from Cape Bathurst” to eat dinner. They 
returned to the same bay about midnight on 10 Au- 
gust, after having been turned back by the pack ice at 
Cape Bathurst, and remained camped there until 15 
August while they waited to see if ice conditions at 
the Cape might improve. This places Pullen within 
one nautical mile of Richardson’s (1828) estimate of 
his position when I believe he made his first collection 
of B. pilosa, and at the perfect season to find the plant 
in fruit. He describes the small bay as being “the only 
place where we could conveniently land,” which sug- 
gests that it is likely the same place Richardson had 
chosen to land for breakfast in 1826 when he discov- 
ered the plant. In addition, Pullen’s steersman was Neil 
McLeod, a Hudson’s Bay Company employee who had 
also accompanied Richardson in 1848. It is probable 
that McLeod would have mentioned Richardson’s 
interest in the plant to Pullen and perhaps inspired 
him to make his own collection during the four days 
they were camped at the site. 

If Richardson’s and Pullen’s B. pilosa collections 
did come from the same location on Cape Bathurst 


Peninsula, the position of that location would most 
likely be between about 70° 27' and 70° 28' north lat- 
itude along the western coast of the peninsula (Figure 
3). Or perhaps slightly farther north if their estimates 
of the distance to Cape Bathurst were based on the 
distance they actually traveled along the coastline 
rather than the direct straight-line distance. This section 
of coastline should be searched carefully for extant 
populations of B. pilosa. 

While the rediscovery of B. pilosa would of itself 
be interesting botanically and historically, the avail- 
ability of living plants for study would also be tremen- 
dously valuable to our understanding of Braya. The 
evolution of the genus is intimately tied to hybridiza- 
tion and polyploidy (Harris 1985), but that phyloge- 
netic history is obscured by the fact that diploidy has 
never been reported in Braya. There is some evidence, 
however, that B. pilosa may be diploid and, therefore, 
could have played a critical role in the evolutionary 
history of Braya. Ornduff (1969) listed several attrib- 
utes in flowering plants that are correlated with xeno- 
gamy and a diploid chromosome number. Among them 
are rotate corollas with large petals, exserted styles, 
scented flowers, abortive fruits, and a narrow distribu- 
tion. Braya pilosa has the largest petals and longest 
styles of any Braya, it is the only species in the genus 
with scented flowers, it appears to have a fairly high 
frequency of abortive silicles, and it has a very narrow 


2004 HARRIS: PILOSE BRAYA: AN ENDEMIC OF ARCTIC CANADA 


SBP) 


FIGURE 4: Photograph of Braya pilosa specimen collected on 24 July 2004 (J. G. Harris and D. L. Taylor 3644). 


556 


distribution. Large, rotate flowers, allogamy, abortive 
fruits, and narrow distributions also have been associ- 
ated with lower ploidy levels in B. humilis (Harris 1985), 
though these are tetraploids rather than diploids. 

Pollen measurements also provide evidence that B. 
pilosa may be diploid. Rollins (1953) and Bécher (1956) 
noted a correlation of ploidy level and pollen size in 
B. humilis, with populations of lower ploidy level 
having somewhat smaller pollen grains than those of 
higher ploidy level. Pollen sizes in B. pilosa, B. thorild- 
wulffii, and B. glabella show a similar correlation. Mea- 
surements (Harris 1985) indicate that B. pilosa has 
significantly smaller pollen grains than the tetraploid 
B. thorild-wulffii, with the largest pollen grains being 
found in the octoploid B. glabella. 

If B. pilosa is indeed diploid, its extremely limited 
distribution may be linked to its ploidy level. Favarger 
(1961) and Johnson and Packer (1967) found a corre- 
lation between plant ploidy level and habitat stability. 
Diploids were more likely to be found on stable habi- 
tats, while polyploids were more frequent on habitats 
with a history of disturbance. Areas that remained 
unglaciated during the Pleistocene, for example, were 
found to have a significantly higher percentage of 
diploids in their floras than areas that were glaciated 
(Favarger 1961; Johnson and Packer 1967). The Cape 
Bathurst Peninsula is known to have been unglaciated 
during the Pleistocene (Prest 1969) and probably served 
as a refugium for plants that moved back onto their 
former ranges as the ice receded. Braya pilosa likely 
weathered the Pleistocene on the relatively stable habi- 
tats available near Cape Bathurst, but it has apparently 
been unable to expand its range in recent times. It may, 
however, have served as a parent species to some of 
the more widespread polyploids in the genus. A like- 
ly candidate is B. thorild-wulffii, a high arctic North 
American endemic that shares B. pilosa’s ovoid- 
ellipsoid fruit shape and fenestrate silicle septae, and 
appears to be most closely related to it. Braya tho- 
rild-wulffii is a tetraploid (2n=28) (Holmen 1952; 
Mulligan 1965, 2002; Bocher 1966; Harris 1985), 
which is the lowest reported ploidy level in the genus 
(some populations of B. humilis are also tetraploid). 
However, Boécher (1966) observed that there appears 
to be only a single large pair of chromosomes in B. 
thorild-wulffii, so it is probably not simply an autote- 
traploid from a single diploid parent. 

DNA sequence data (Harris, unpublished data) sug- 
gest that B. thorild-wulffii may have served as a parent 
to other Braya species of higher ploidy level. If DNA 
of B. pilosa were available, and if the species is indeed 
a parent to B. thorild-wulffii, it could provide signifi- 
cant insight into phylogenetic relationships within Braya 
and between Braya and other genera in the Cruciferae. 


Acknowledgments 
I am indebted to C. Stuart Houston for his help with 
historical questions about the second Franklin expe- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


dition, Ihsan Al-Shehbaz for his helpful discussions 
about the taxonomic status of Braya pilosa, and Alvan 
Bregman of the Rare Book Collections Library at Uni- 
versity of Illinois at Urbana-Champaign for providing 
me with transcripts of Richardson’s annotations to his 
personal copy of Flora Boreali-Americana. 1 thank 
Sam Rushforth, Renée Van Buren, Kimball Harper, 
and JaNae Brown Haas for insightful comments on 
previous drafts of this paper and Melinda Woolf Har- 
ris for her illustrations of Braya pilosa. Finally, | am 
indebted to two anonymous reviewers who provided 
valuable comments and suggestions. 


Addendum 

On 24 July 2004, Daniel L. Taylor and I briefly visit- 
ed Richardson’s presumed type locality of Braya pilosa 
near Cape Bathurst. We discovered several hundred 
B. pilosa individuals growing along the coastline on 
sandy, calcareous soils in an area heavily used by Cari- 
bou (Ranifer taranclus). The plants were limited to 
small, bare patches of soil disturbed by Caribou hoofs. 

The plants on Cape Bathurst Peninsula (Figure 4) 
are perfect matches for Richardson’s and Pullen’s 
19" Century B. pilosa collections (Figure 2). Speci- 
mens (J. G. Harris and D. L. Taylor 3644) are deposit- 
ed in the Utah Valley State College Herbarium (UVSC), 
with duplicate material distributed to the following 
herbaria: Agriculture and Agri-Food Canada (DAO), 
Canadian Museum of Nature (CAN), Missouri Botan- 
ical Garden (MO), New York Botanical Garden (NY). 
DNA sequencing and cytological studies are currently 
underway (Harris, unpublished data). 


Documents Cited (marked * in text) 

Mulligan, G. A. 2000. Key to the Brassicaceae (Cruciferae) 
of Canada and Alaska. Electronic publication on the inter- 
net site, http://members.rogers.com/mulligan4520/key/ 

Murray, D. F. 1983. in sched., Mouth of River Mackenzie, 
Richardson s.n., National Herbarium of Canada (CAN) 
64149, Ottawa, Ontario. 

Scott, P. J., S. G. Aiken, and M. J. Dallwitz. 2001. Brassi- 
caceae of the Canadian Arctic Archipelago: descriptions, 
illustrations, identification, and information retrieval. Ver- 
sion: 29" June 2001. Electronic publication on the internet 
site, http://www.mun.ca/biology/delta/arcticf/ 

Warwick, S. I., A. Francis, and G. A. Mulligan. 2000. Bras- 
sicaceae of Canada. Agriculture and Agri-Food Canada, 
Ottawa. Electronic publication on the internet site, http:// 
sis.agr.gc.ca/brd/brass/ 


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ska Vetenskapsakademiens Handlingar. Fjarde Serien. Band 
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ford, California. 

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ralized mustards, Brassicaceae (Cruciferae). Canadian 
Field-Naturalist 116(4): 611-622. 

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Received 20 October 2003 
Accepted 6 December 2004 


New Records of Vascular Plants in the Yukon Territory VI 


WILLIAM J. Copy!, CATHERINE E. KENNEDY”, BRUCE BENNETT’, and PHIL CASWELL? 


‘Biodiversity, National Program on Environmental Health, Agriculture and Agri-Food Canada, Wm. Saunders Building (49), 
Central Experimental Farm, Ottawa, Ontario KIA 0C6 Canada 

*Department of Environment, Government of the Yukon, Box 2703, Whitehorse, Yukon Y1A 2C6 Canada 

3P. O. Box 91, Roxbury, New York 12474 USA 


Cody, William J., Catherine E. Kennedy, Bruce Bennett, and Phil Caswell. 2004. New records of vascular plants in the Yukon 
Territory VI. Canadian Field-Naturalist 118(4): 558-578. 


Based on field reconnaissance mainly in 2002 in the southern part of the Yukon and particularly in and adjacent to Kluane National 
Park, information is provided on geographically significant plant occurrences. Six native taxa, Atriplex alaskensis, Claytonia 
megarrhiza, Corispermum ochotense vat. alaskanum, Oxytropis arctica, Polemonium acutiflorum forma lacteum and Polemo- 
nium boreale forma albiflorum, and four introduced taxa. Arabis caucasica, Camelina sativa, Senecio eremophilus, and Setaria 
viridis are reported new to the known flora of the Yukon Territory. Significant range extensions for 158 native and 21 introduced 
taxa are included. Parrya arctica, Armoracia rusticana, Atriplex patula and Papaver nudicaule ssp. nudicaule are excluded 


from the Yukon flora. 


Key Words: Vascular plants, Yukon Territory, flora, new records, range extensions, phytogeography. 


Since the writing of New Records of Vascular Plants 
in the Yukon Territory V (Cody et al. 2003), a consider- 
able number of plant specimens have been submitted 
to Cody for identification and confirmation. The major 
submissions include the following locations and col- 
lectors: (1) Kluane National Park and vicinity by Phil 
Caswell for the National Park Service as part of on- 
going botanical inventories; (2) W. J. Cody and Cath- 
erine Kennedy spent four days collecting specimens in 
the Fort Selkirk area assessing the flora of that region; 
(3) W. J. Cody and Margaret Cody studied changed 
vegetation along the southern highways; (4) Bruce Ben- 
nett conducted studies adjacent to the North Canol 
Road, assisting with rare plant surveys for Yukon- 
Charley Rivers National Park and interesting areas 
while boating down the Yukon River; (5) Rhonda Rosie 
visited sites adjacent to Nisling Lake and Morris Lake 
for the Yukon Territory Government and Parks Branch; 
(6) Stu Withers visited former mine sites at Atlin Lake 
and Deadman Creek in connection with reclamation of 
vegetation and soil stabilization; (7) with his continuing 
interest in vegetation habitats Greg Brunner collected 
plant specimens from areas adjacent to the Klondike 
River and Dawson City. 

This paper serves to further update the Flora of the 
Yukon Territory (Cody 1996) and Flora of the Yukon 
Territory, Second Edition (Cody 2000) along with other 
records recently published (Cody et al. 1998, 2000, 
2001, 2002, 2003). The floristic information presented 
earlier and updated here is essential for biological re- 
search and ongoing work relating to agriculture, fores- 
try, sustainable resource management and wildlife 
management. With additions of six native and four 
introduced species reported here, the flora now includes 
1181 species. The new native species are all rare (as 
defined by Douglas et al. 1981). 


The taxa addressed in the body of this paper appear 
in a synoptic list by Yukon status in alphabetical order. 
The taxa are then discussed in taxonomic order, as pre- 
sented in the Flora of the Yukon Territory with citation 
of specimens and other pertinent information. Common 
names follow Cody (1996) and Douglas et al. (1998- 
2001). 


Synoptic List by Yukon Status 

Errata in New Records: (5) 

Carex stylosa 

Douglasia arctica 

Poa nemoralis revised to Poa interior 

Armoracia rusticana revised to Rorippa barbareifolia 
Papaver nudicaule ssp. nudicaule revised to Papaver croceum 


Native Taxa New to the Yukon Territory: (6) 
Atriplex alaskensis 

Claytonia megarrhiza 

Corispermum ochotense var. alaskanum 

Oxytropis arctica 

Polemonium acutiflorum forma lacteum 

Polemonium boreale forma albiflorum 


Introduced Taxa New to the Yukon Territory: (4) 
Arabis caucasica 

Camelina sativa 

Senecio eremophilus 

Setaria viridis 


Range Extensions of Native Taxa within the Yukon 
Territory: (158) 

Agrostis filiculmis 

Allium schoenoprasum ssp. sibiricum 

Alyssum obovatum 

Amelanchier alnifolia 

Arabis holboellii var. retrofracta 

Arabis holboellii var. secunda 

Aster alpinus ssp. vierhapperi 

Aster falcatus 


558 


Atriplex subspicata 

Bromus ciliatus 

Callitriche anceps 

Cardamine bellidifolia 

Carex albo-nigra 

Carex athrostachya 

Carex atrosquama 

Carex capillaris ssp. chlorostachys 
Carex crawfordii 

Carex eleusinoides 

Carex foenea 

Carex limosa 

Carex loliacea 

Carex obtusata 

Carex pachystachya 

Carex pellita 

Carex petasata 

Carex praegracilis 

Carex rossii 

Carex supina ssp. spaniocarpa 
Carex sychnocephala 

Carex viridula 

Cassiope tetragona ssp. saximontana 
Cerastium arvense 
Chamaerhodos erecta ssp. nuttallii 
Chenopodium capitatum 

Cicuta maculata var. angustifolia 
Cryptogramma crispa vat. acrostichoides 
Deschampsia brevifolia 
Descurainia incisa vat. incisa 
Douglasia alaskana 

Draba aurea 

Draba corymbosa 

Draba crassifolia 

Draba glabella 

Draba nemorosa 

Draba ogilviensis 

Draba palanderiana 

Draba porsildii 

Draba stenoloba 

Dryas hookeriana 

Eleocharis uniglumis 

Elymus alaskanus ssp. alaskanus 
Elymus glaucus 

Elymus macrourus 

Elymus spicatus 

Elymus trachycaulus ssp. andinus 
Erigeron caespitosus 

Erigeron hyperboreus 
Eriophorum brachyantherum 
Eritrichium aretioides 
Eritrichium chamissonis 
Euphrasia subarctica 

Festuca brachyphylla 

Festuca richardsonii 

Festuca saximontana 
Gentianella tenella 

Geum aleppicum ssp. strictum 
Geum rossii 

Glyceria striata var. stricta 
Gymnocarpium jessoense ssp. parvulum 
Harrimanella stellariana 
Hordeum brachyantherum 

Tris setosa ssp. interior 

Isoetes echinospora 


2004 Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON VI 559 


Isoetes ?maritima 

Juncus bufonius 

Juncus filiformis 

Kobresia simpliciuscula 

Koeleria macrantha 

Lepidium densiflorum var. densiflorum 
Lupinus nootkatensis 

Luzula arcuata ssp. unalaschkensis 
Maianthemum canadense ssp. interius 
Mertensia paniculata var. alaskana 
Mimulus guttatus 

Minuartia dawsonensis 

Minuartia elegans 

Minuartia macrocarpa 

Minuartia yukonensis 

Montia fontana 

Myriophyllum verticillatum 
Nuphar polysepalum 

Orobanche fasciculata 

Oxytropis campestris ssp. varians 
Pedicularis oederi 

Phacelia mollis 

Phlox hoodii 

Phyllodoce glanduliflora 
Pinguicula villosa 

Pinus contorta ssp. latifolia 
Plantago canescens 

Plantago maritima 

Poa interior 

Poa leptocoma 

Poa secunda 

Podistera macounii 

Polygonum lapathifolium 
Potamogeton alpinus ssp. tenuifolius 
Potamogeton berchtoldii 
Potamogeton gramineus 
Potamogeton pectinatus 
Potamogeton praelongus 
Potamogeton subsibiricus 
Potamogeton zosteriformis 
Potentilla arguta var. convallaria 
Primula eximia 

Puccinellia interior 

Pulsatilla ludoviciana 

Pyrola minor 

Ranunculus aquatilis var. subrigidus 
Ranunculus occidentalis var. brevistylis 
Rhinanathus minor ssp. borealis 
Ribes oxyacanthoides ssp. oxyacanthoides 
Rorippa barbareifolia 

Rosa woodsii 

Salix alaxensis ssp. longistylis 
Salix glauca vat. acutifolia 

Salix pseudomyrsinites 

Salix rotundifolia ssp. dodgeana 
Sambucus racemosa ssp. pubens 
Saxifraga caespitosa 

Saxifraga eschscholtzii 

Saxifraga foliolosa 

Saxifraga nelsoniana ssp. pacifica 
Saxifraga rufopilosa 

Saxifraga tricuspidata 

Scirpus rollandii 

Silene involucrata ssp. tenella 
Silene williamsii 


560 


Solidago canadensis vat. salebrosa 
Spiranthes romanzoffiana 
Stellaria umbellata 

Stipa comata 

Stipa nelsonii ssp. dorei 

Stipa richardsonii 

Subularia aquatica ssp. americana 
Tofieldia coccinea 

Trientalis europaea 

Triglochin palustre 

Typha latifolia 

Urtica dioica ssp. gracilis 
Utricularia minor 

Vaccinium membranaceum 
Vahlodea atropurpurea 

Valeriana sitchensis 

Vicia americana 

Viola renifolia var. brainerdii 
Woodsia alpina 

Woodsia ilvensis 


Range Extensions of Introduced Taxa within the 
Yukon Territory: (21) 

Agropyron pectiniforme 

Agropyron sibiricum 

Agrostis gigantea 

Avena sativa 

Brassica rapa 

Bromus inermis 

Capsella bursa-pastoris 

Elytrigia intermedia 

Galeopsis tetrahit ssp. bifida 

Lepidium densiflorum var. macrocarpum 
Myosotis scorpioides 

Papaver croceum 

Poa annua 

Rheum rhabarbarum 

Rumex crispus 

Silene vulgaris 

Sinapis arvensis 

Sonchus arvensis ssp. uliginosus 
Tanacetum vulgare 

Thlaspi arvense 

Trifolium pratense 

Comments on Native Taxa in the Yukon Flora: (3) 
Carex stylosa 

Douglasia arctica 

Isoetes maritima 


Deletions of Native Taxa from the Yukon Flora: (1) 
Parrya arctica 


Deletions of Introduced Taxa from the Yukon Flora: 
(3) 

Armoracia rusticana 

Atriplex patula 

Papaver nudicaule ssp. nudicaule 


Annotated Species List 

ISOETACEAE 

Isoetes echinospora Dut., Bristle-like Quillwort — 
YUKON: On stony shore of small pond south of Morris 
Lake, 60°20°N 131°40°W, R. Rosie 2030, 8 Sept. 1999 
(DAO); washed up on lakeshore, northeast shore of 
Nisling Lake, 62°02’00"N_ 137°58’30"W, R. Rosie 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


02-821, 12 Aug. 2002 (DAO) (determined by D. F. 
Brunton). 

The first specimen cited above is an extension of the 
known range in the Territory (Cody 1996) of about 180 kilo- 
meters southwest of the vicinity of Francis Lake. The second 
specimen is from a site about 500 kilometers northwest of 
Frances Lake and about 430 kilometers northwest of the 
Morris Lake site. 


Isoetes ?maritima Underw., Maritime Quillwort — 
YUKON: washed ashore, WNW side of Trout Lake, 
68°49°27"N 138°45'8"W, J. M. Line 2000-72, 28 July 
2000 (DAO) (determined by D. F. Brunton & D. M. 
Britton). 

The fragmented specimens found washed up along the 
shore of the lake possessed only immature female spores 
which were mostly densely tuberculate but could not defi- 
nitely be determined as /. maritima. If this could be verified 
by more mature specimens from Trout Lake it would be only 
the third known location in the Territory and a range exten- 
sion of about 260 kilometers northwest of Vittrewka Lake 
(Cody et al. 2001). 


Isoetes maritima Underw., Maritime Quillwort — It 
has been brought to our attention by Lori Schroeder 
(Environment Yukon) that specimen data according 
to Greg Brunner’s field notebook for his collection 
#39-99 which was published in Cody et al. (2001) 
should be: Peel River Wetland Study Turner Lake, 
66°11'N 134°14'W, 2 July 1999. 


PTERIDACEAE 
Cryptogramma crispa (L.) R. Br. var. acrostichoides 
(R. Br.) C. B. Clarke, Mountain-parsley — YUKON: 
Kluane National Park, south-facing scree slope, 
60°06.019'N 137°08.532'W, P. Caswell 026, 6 June 
2002 (DAO). 

This rare species which was previously known from only 
six localities in the Territory is now known from two sites in 
southern Kluane National Park. 


ASPIDIACEAE 
Gymnocarpium jessoense (Koidz.) Koidz. ssp. parvu- 
lum Sarvela, Nahanni Oak Fern — YUKON: dry rock out- 
crop, lower Beaver River, 60°09'09"N 124°55'16"W, 
B. Bennett 97-518, 19 Aug. 1997 (B. Bennett Herbar- 
ium, photo DAO). 

The specimen cited above of this rare plant in the 
Territory is an extension of the known range of about 


300 kilometers from a site in the vicinity of Francis 
Lake (Cody 1996). 


Woodsia alpina (Bolton) S. F. Gray, Northern Woodsia — 
YUKON: damp organic soil in rock crevice, Cache Lake, 
Kluane National Park, 61°12.586'N 139°03.671'W, P. 
Caswell 575, 24 July 2002 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 45 kilometers southwest of a 
site at the north end of Kluane Lake and is new to the flora 
of Kluane National Park. 


Woodsia ilvensis (L.) R. Br., Rusty Woodsia — YUKON: 
steep south-facing bluffs along Klondike River, Dawson 
Area, | mile north of 64°02'N 137°41'30"W, G. 
Brunner 435, 2 June 2001 (DAO); dry rocky outcrop, 


2004 


S Fork bluffs, Klondike River, 64°O1'N 138°9'W, G. 
Brunner 443-01, 14 July 2001 (DAO). 

Douglas et al. (1981) considered this species rare in the 
Territory. The specimens cited above are from sites about 
60 kilometers southwest of a site adjacent to the Dempster 
Highway mapped by Cody (1996). 


PINACEAE 

Pinus contorta Doug}. ex Loud. ssp. latifolia Engelm. 
ex S. Wats., Lodgepole Pine — YUKON: roadside, North 
Canol Road KM 284, 62°13'02"N 131°46'10"W, B. 
Bennett 02-723, 22 Sept. 2002 (DAO); roadside across 
road from large truck dump, North Canol Road Km 
380, 62°50'03"N 130°56'24"W, B. Bennett 02-724, 
21 Sept. 2002 (DAO). 

The specimens cited above are an extension of the known 
range in the Territory of about 175 kilometers northeast of a 
site adjacent to the South Canol Road. The second specimen 
was from a site adjacent to an old house and may have been 
planted. 


TYPHACEAE 

Typha latifolia L., Common Cattail — YUKON: border 
of small lake below swimming pool, Takhini Hot 
Spring, 60°52'40.7"N 135°21'30.6"W, Cody & Cody 
37692, 10 Aug. 2001 (DAO); same locality (fruiting) 
Cody 35311, 4 Aug. 2002 (DAO). 

Douglas et al. (1981) considered this species rare in the 
Territory. Cody (1996) knew it only from the vicinity of 
Mayo. Cody et al. (2000, 2001) added additional sites in the 
extreme southeast and in the vicinity of Faro. The specimen 
cited above which was collected just west of Whitehorse is 
now from the fourth site in the Territory. 


POTAMOGETONACEAE 

Potamogeton alpinus Balbis ssp. tenuifolius (Raf.) 
Hultén — YUKON: plant fragments washed ashore, Trout 
Lake, 68°50'N 138°45'W, J. M. Line 2000-9, 29 July 
2000 (DAO). 

The specimen cited above is the northernmost yet found 
in the Territory. It is an extension of the known range (Cody 
1996) of about 85 kilometers NE of a site north of latitude 
68°N. 


Potamogeton berchtoldii Fieb. — YUKON: shallow 
water of small lake, west side of Dempster Highway 
Km 93, 64°39'01"N 138°23'01"W, Cody & Cody 
38060, 27 July 2002 (DAO). 

The specimen cited above is from a site about 80 kilo- 
meters northeast of Dawson City. To the north a single site 
was previously known from adjacent to the Peel River (Cody 
1996). 


Potamogeton gramineus L., Variable-leaved Pondweed 
— YUKON: slough off Yukon River with old sunken 
dredge, 61°56'35"N 135°12'17"W, B. Bennett 02-299, 
7 Aug. 2002 (DAO). 

The nearest site to that listed above known to Cody (1996) 
is about 75 kilometers to the north. 


Potamogeton pectinatus L., Sago Pondweed — YUKON: 
shallow backwater, Dezadeash River, Kluane National 
Park, 60°45.683'N 137°36.010'W, P. Caswell S05, 16 
Aug. 2002 (DAO); locally common in mucky alka- 


Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON VI 


56] 


line pond, Mile 985 Alaska Highway, 60°48'N 
136°45'W, Calder & Kukkonen 28277, 14 Aug. 1960 
(DAO). 

The specimens cited above extend the known range in the 

Territory about 120 kilometers west of the vicinity of White- 
horse. The Caswell specimen is the first known from Kluane 
National Park. 
Potamogeton praelongus Wulf., White-stemmed 
Pondweed — YUKON: Enoch Lake, Old Crow Flats, 
68°05'00"N_ 140°09'50"W, B. Bennett 95-413, 10 
Aug. 1995 (B. Bennett Herbarium, photo DAO); 
plant fragments washed ashore, Trout Lake, 68°50'N 
138°45'W, J. M. Line 2000-140, 28 July 2000 (DAO); 
Nordenskiold River wetland pond, M. Dennington, 
Nord #23, 19 July 1984 (B. Bennett Herbarium, photo 
DAO). 

Douglas et al. (1981) considered this species rare in the 
Yukon Territory based on specimens from only two locations. 
Cody (1996) knew it from five locations. Cody et al. (2000) 
added three locations. The first specimen cited above is a 
northwestward extension of about 40 kilometers from the 
vicinity of Old Crow. The second specimen from about 115 
kilometers NNE of Old Crow is the northernmost yet found 
in the Territory. The third specimen is an extension of the 
known range in the south of about 125 kilometers northwest 
of a site just north of Whitehorse. 


Potamogeton subsibiricus Hagstr. - YUKON: growing 
in about 3 feet of water, lake west of Dempster Rd. 
near mile 65, R. T: Porsild 317, 25 July 1966 (AKA, 
photo DAO) (determined by A. Batten). 

Cody (1996) knew this rare species in the Territory from 
a single collection from adjacent to the North Canol Road 
about 450 kilometers to the southeast of the locality cited 
above. 


Potamogeton zosteriformis Fern., Flatstemmed Pond- 
weed — YUKON: in water by outlet of pond, Pumphouse 
Pond, 3 km S of Alaska Highway, 60°43'31.3"N 
135°09'58.3"W, W. J. Cody 38290, 3 Aug. 2002 
(DAO). 

Cody (1996) knew this rare species in the Territory from 
only two localities, one southeast of Haines Junction and one 
just north of 64°N. Cody et al. (1998, 2000 and 2001) added 
two sites in the south and one north of 68°N. 


SCHEUCHZERIACEAE 
Triglochin palustre L., Marsh Arrow-grass — YUKON: 
silty river bank with Carex utriculata, C. aquatilis 
and Juncus alpinoarticulatus, Selwyn, Yukon River, 
62°48'06"N 138°15'29"W, B. Bennett 02-294, 9 Aug. 
2002 (DAO). 

The specimen cited above is between sites adjacent to 
the Alaska Highway, Klondike Highway and Dawson City. 


POACEAE (GRAMINEAE) 

Agropyron pectiniforme R. & S., Crested Wheat 
Grass — YUKON: lake shore, Earn Lake, 62°.9'08.3"N 
134°23'07.6"W, G. Brunner 557-01, 5 Aug. 2001 
(DAO); steep stony slope on east side of highway, 
Haines Highway, 60°6'3.2"N 136°54'51"W, Cody & 
Cody 37911, 21 July 2002 (DAO). 


562 


The first specimen of this introduced species cited above 
is from a site between the northernmost mapped by Cody 
(1996) in the vicinity of Mayo and a site adjacent to Faro. 
The second specimen cited above is an extension of the 
known range in the Territory of about 85 kilometers south- 
east of a site in the vicinity of Haines Junction. 


Agropyron sibiricum (Willd.) P. Beauv. — YUKON: top of 
bank overlooking road into gravel pit south of Alaska 
Highway, 60°49'58"N 135°45'49"W, Cody & Cody 
38210, 1 Aug. 2002 (DAO). 

Cody (1996) knew this introduced species in the Territory 
only from two locations, Whitehorse and Carmacks. A third 
site was reported from the vicinity of Ross River (Cody et al. 
2003). The specimen cited above is from a site about 40 kilo- 
meters west of Whitehorse. 


Agrostis filiculmis M. E. Jones — YUKON: open area — 
abandoned placer mine, Mechanic Creek, 62°20.1'N 
137°21.5'W, S. Withers SWOI-062, 6 July 2001 (DAO). 
The specimen cited above is only the second known in the 
Territory (Cody 1996) and is from a site about 200 kilometers 
southeast of the first site in the vicinity of Dawson City. 


Agrostis gigantea Roth., Creeping Bent Grass — 
YUKON: Spruce Hill Park, border of playground at 
head of Engelman Drive, Whitehorse, 60°36'40.3"N 
134°55'08.8"W, W. J. Cody 37743, 14 July 2002 
(DAO). 

Cody (1996) knew this introduced species in the Territory 
from only two localities, Dawson City and Carmacks. An 
additional collection from a mine site east of Dawson City 
was reported by Cody et al. (2003). The specimen cited above 
is from only the fourth known locality in the Territory. 


Avena sativa L., Oats — YUKON: scattered in grass area 
around Beringia Museum, Whitehorse, 60°42'36"N 
135°4'45"W, Cody & Cody 38105, 30 July 2002 
(DAO). 

Cody (1996) knew this introduced species from only wide- 
ly separated sites in the Territory, vicinity of Mayo, middle 
of North Canol Road and east of Watson Lake. 


Bromus ciliatus L. — YUKON: in moist moss on moun- 
tain slope above treeline on east side of Dempster 
Highway about 9 kilometers north of Tombstone Park 
Campsite, 64°34'49"N 138°15'30"W, Cody & Cody 
38047, 26 July 2002 (DAO). 

The specimen cited above is at the northern limit of this 
species in the Territory (Cody 1996). The nearest sites previ- 
ously mapped were about 120 kilometers to the west adjacent 
to the Yukon River and about 150 kilometers to the southeast 
in the vicinity of Stewart Crossing. This species was con- 
sidered rare in the Territory by Douglas et al. (1981). 


Bromus inermis Leyss., Smooth Brome — YUKON: 
sand beach, Kusawa Lake Campground, 60°5'10"N 
136°8'42"W, Cody & Cody 38144, 31 July 2002 
(DAO); gravel ditch between road and steep gravel 
slope, Haines Highway, 60°6'3.3"N 136°54'51"W, 
Cody & Cody 37912, 21 July 2002 (DAO). 

The specimens cited above of this introduced species 
which are from sites about 100 kilometers southeast of Haines 
Junction and southwest of Whitehorse, are the most south- 
western yet found in the Territory. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Deschampsia brevifolia R. Br. — YUKON: flats 7 miles 
east of Tagish Bridge, 60°19'5"N 134°10'17"W, Cody 
& Cody 37940, 23 July 2002 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory (Cody 1996) of about 90 kilometers 
southeast of a site west of Whitehorse. 


Elymus alaskanus (Scribn. & Merr.) A. Léve ssp. 
alaskanus — YUKON: in sandy soil of river bank, Nis- 
utlin River Delta National Wildlife Area, 60°11'N 
132°35'W, B. Bennett 98-167, 26 July 1998 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 180 kilometers south of a site 
adjacent to the northern South Canol Road. 


Elymus glaucus Buckl., Western Rye Grass — YUKON: 
roadstop in disturbed gravelly soil, Atlin Road just 
north of BC border, 60°00'09"N 133°47'42.1"W, 
Cody & Cody 37547, 28 July 2001 (DAO) 

Douglas et al. (1981) considered this species rare in the 
Territory. The specimen cited above 1s an extension of about 
115 kilometers southeast of a site adjacent to Whitehorse 
reported by Cody et al. (2002). 


Elymus macrourus (Turcez.) Tzvelev — YUKON: grow- 
ing in loose sand on riverbar, Wind River, camp #6, 
65°40.46'N 135°11.76'W, B. Bennett 00-853, 7 July 
2000 (B. Bennett Herbarium, photo DAO); exposed 
sandy lakeshore, north end of Atlin Lake, 60°01.5'N 
133°50.4'W, S. Withers SWO1-045, 30 June 2001 
(DAO). 

The first specimen of this Amphi-Beringian species cited 
above is the first known from between latitudes 64°N and 
66°N east of the Dempster Highway. The second specimen 
is an extension of the known range in the Territory of about 
200 kilometers southwest of a site east of Haines Junction. 


Elymus spicatus (Pursh) Gould — YUKON: common at 
base of large talus slope, Yukon River, Brittania Creek, 
62°52'28"N 138°42'56"W, B. Bennett 02-641, 10 Aug. 
2002 (DAO). 

Cody (1996) knew this species in the southwest of the Ter- 
ritory and disjunct to the extreme northwest. The specimen 
cited above is an extension of the known range in the 
Territory of about 85 kilometers west of longitude 137°W. 


Elymus trachycaulus (Link) Gould ex Shinners ssp. 
andinus (Scribn. & Smith) A. & D. Léve — YUKON: 
gravel roadside, Top of the World Highway Km 16, 
64°6'6"N 139°38'41"W, Cody & Cody 38035, 38037, 
26 July 2002 (DAO); gravel roadside, Top of the World 
Highway Km 102, 64°05'33"N 140°55'22"W, Cody 
& Cody 38003, 26 July 2002 (DAO). 

The specimens cited above extend the known range in the 
Territory to the west about 140 kilometers from a site east of 
the Dempster Highway. 


Festuca brachyphylla Schultes & Schultes f., Short- 
leaf Fescue — YUKON: Site KPL #106, 62°17'38.7"N 
137°48'25"W, G. Brunner 578-01, July 2001 (DAO). 

The specimen cited above is from a site between sites 
mapped by Cody (1996) west of Carmacks and adjacent to 
southern Dempster Highway. 


2004 


Festuca richardsonii Hooker — YUKON: beside test plot, 
| km N of Montague Roadhouse, Km 132 Klondike 
Highway, 61°48'35"N_ 136°7'52"W, Cody & Cody 
38082, 28 July 2002 (DAO). 

This species is frequent in the southern part of the Terri- 
tory west of longitude 137°W and north of latitude 64°N 
but has not previously been found adjacent to the Klondike 
Highway. 


Festuca saximontana Rydb., Rocky Mountain Fescue 
— YUKON: gravel beside road, Alaska Highway 8 km 
west of White River, 62°3'15.6"N 140°38'20.5"W, 
Cody & Cody 37886, 37887, 20 July 2002 (DAO). 

The specimens cited above are an extension of the known 
range in the Territory of about 100 kilometers northwest of 
a site just northwest of Kluane Lake. 


Glyceria striata (Lam.) Hitche. var. stricta (Scribn.) 
Fern., Fowl Marina Grass — YUKON: moist muddy 
organic soil, along road from Dalton Post to Wade 
Lakes, 60°07.765'N 137°04.105'W, P. Caswell 650, 4 
Aug. 2002 (DAO). 

This plant was considered rare in the Territory by Douglas 
et al. (1981). Cody (1996) knew it only in the southeast as 
far west as just west of Watson Lake. The specimen cited 
above from between the Haines Highway and Kluane Na- 
tional Park is from a site about 450 kilometers to the west. 
It is however known from British Columbia. 


Hordeum brachyantherum Nevski — YUKON: disturbed 
area in townsite, Big Salmon Village at confluence of 
Yukon River, 61°53'N 134°55'W, B. Bennett 02-292, 
7 Aug. 2002 (DAO). 

Cody (1996) considered this species to be introduced at 
Dawson City but native and rare at Carcross. Sites in the 
vicinity of Whitehorse (Cody et al. 2001) were considered 
to be introduced. The specimen cited above was probably 
also introduced. 


Koeleria macrantha (Ledeb.) Schultes, June Grass — 
YUKON: south aspect-xeric-sage/graminoid-forbs of 
low compact growth, Site YPC103 near confluence of 
Pelly and Yukon rivers, 62°48'16"N 137°19'35.9"W, 
G. Brunner 536-01, 3 Aug. 2001 (DAO) (determined 
by S. Darbyshire); gravel pit just north of the Alaska 
Highway, Silver City Road, 61°01'22"N 138°19'52"W, 
B. Bennett 01-005, 29 Apr. 2001 (DAO); lower slope 
south-facing, dry, Five Fingers Coal Mine, Yukon River, 
62°12'21"N 136°20'10"W, B. Bennett 02-295, 8 Aug. 
2002 (DAO). 

Douglas et al. (1981) considered this species rare in the Ter- 
ritory. The first specimen cited above was collected just south- 
west of the Pelly River Ranch at 62°50'20"N 137°11'40"W 
by M. Johansen in 1989 (DAO) (Cody 1996); the second 
specimen is an extension of about 60 kilometers northwest 
of a site adjacent to Haines Junction; the third specimen is 
from a site about 40 kilometers southeast of a site near Minto 
(Cody 1996). 


Poa annua L., Annual Blue Grass — YUKON: vicinity of 
campground, Rancheria, at mile 710 Alaska Highway 
(60°0S'N 130°36'W), S. L. Welsh & G. Moore 7539, 
29 June 1968 (AKA, photo DAO) (determined by R. 
J. Soreng). 


Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON VI 


563 


Cody (1996) knew this introduced species from only five 
sites in the Territory. The specimen cited above is about 100 
kilometers west of a site adjacent to the southern Canol Road. 
Poa interior Rydb. (P. nemoralis sensu Cody 1996) — 
YUKON: in subalpine meadow, Onion Lake, ca. 46 mi 
S of Haines Junction, G. W. & G. G. Douglas 7089, 
12 Aug. 1973 (AKA, photo DAO) (determined by R. 
J. Soreng); common on steep, open, west-facing prairie 
slopes, Conglomerate Mountain on Dawson-White- 
horse road, 61°38'N 135°53'W, Calder & Gillett 
25770, 22 June 1960 (DAO); common on south-facing, 
prairie slope with scattered aspen, Mile 30 on road to 
Dawson from Stewart Crossing, 63°33'N 137°25'W, 
Calder & Gillett 25037, 4 June 1960 (DAO); grown on 
open southern slope, Pelly Ranch (ca. 62°49'N 
136°34'W), J. Y. Tsukamoto s.n., 23 July 1960 (DAO); 
silt bluff, south-facing, on Porcupine River, 1.5 km 
upstream from junction between Rat Indian Creek 
and Porcupine River, 67°34'N 138°21'W L. Cwyner 
977, 22 July 1976 (DAO) (determined by M. Bark- 
worth); dry open exposed gravel ridge overlooking delta, 
west side of river on ridge campsite, Lower Blow 
River Delta, 69°53'N 137°10'W, H. L. Dickson & D. 
L. Allen 5324, 26 July 1982 (DAO); open meadow- 
abandoned beaver pond, Deadman Creek, 60°20.2'N 
133°03.5'W, S. Withers SWO1-126, 16 July 2001 (DAO) 
(determined by S. Darbyshire). ; 

Yukon specimens previously mapped as Poa nemoralis 
by Cody (1996) have been revised to P. interior. In addition, 
the specimens cited above which were inadvertently missed 
have now been mapped. (See new map Figure 1). 


Poa interior 


FiGurE 1. New distribution map for Poa interior in the Yukon 
Territory. 


564 


Poa leptocoma Trin. — YUKON: steep gravelly slope 
adjacent to Alaska Highway 3 km east of Snag Junc- 
tion, 62°12'42.3"N 140°41'33.1"W, Cody & Cody 
37863, 20 July 2002 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 175 kilometers northwest of 
a site in Kluane National Park. 


Poa secunda Presl, Sandberg Bluegrass — YUKON: 
commonly found growing on the base of the bluff and 
occasionally on midslope, Minto Bluff, 62°36'18"N 
136°51'06"W, B. Bennett 02-013, 9 June 2002 (DAO). 

The specimen cited above is an extension of about 175 kilo- 
meters northwest of a site north of Whitehorse (Cody 1996). 
To the north the only other site in the Territory is in the vicin- 
ity of Dawson City. 


Puccinellia interior Th. Sor. — YUKON: gravel roadside, 
Top of the World Highway Km 102, 64°05'33"N 
140°55'22"W, Cody & Cody 38002, 26 July 2002 
(DAO). 

The specimen cited above is an extension of the known 
range in the Territory (Cody 1996) of about 75 kilometers 
west from the vicinity of Dawson City. 


Setaria viridis (L.) Beauv., Green Bristlegrass (Fig- 
ure 2) — YUKON: in crack at edge of Extra Foods park- 
ing lot, downtown Whitehorse, B. Bennett 95-437, 14 
Oct. 1988 (B. Bennett Herbarium, photo DAO). 

The specimen cited above is a new introduction to the 
Flora of the Yukon Territory (Cody 1996). This species which 
is introduced from Eurasia is known across Canada from 
Newfoundland to British Columbia and has been found at 
Fort Simpson in the Northwest Territories. The genus Setaria 
can be separated from the genus Panicum as follows: 


A.  Spikelet surrounded by 1|-many distinct or 
more or less connate bristles, these 
forming aninvolucre eaters ce ee ee Setaria 


B. Spikelet not subtended by bristles 


Stipa comata Trin. & Rupr., Needle-and-thread — 
YUKON: 45° silty sand south-facing slope with Arte- 
mesia frigida and Calamagrostis purpurascens, Dutch 
Bluff, Yukon River, 61°55'N 135°03.99'W, B. Bennett 
02-289, 7 Aug. 2002 (DAO). 

Cody (1996) knew this species in the Territory from only 
four sites south of 63°N. Cody et al. (2003) added an addi- 
tional site just east of Haines Junction. The specimen cited 
above is from a site about 60 kilometers southeast of Car- 
macks. 


Stipa nelsonii Scribn. ssp. dorei Barkworth & Maze 
— YUKON: treeless rocky slope overlooking lake, 
Snafu Lake Camp Site, 60°08'10"N 133°48'23.6"W, 
Cody & Cody 37567, 28 July 2001 (DAO); open area 
adjacent to Populus tremuloides woodland overlook- 
ing lake, 7 km east of Tagish Bridge, 60°19'5"N 
134°10'17"W, Cody & Cody 37938, 23 July 2002 
(DAO). 

Cody (1996) knew this species from only six sites in the 
Territory north to about latitude 61°N. Cody et al. (2001) 
added an additional site in the vicinity of the Carcross Dunes. 


The specimens cited above are from east and southeast of 
the Carcross Dunes site. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


FIGURE 2. Setaria viridis, Green Bristlegrass (drawn by Lee 
Mennell). 


Stipa richardsonii Link (Achnatherum richardsonii 
(Link) Barkw.), Spreading Needlegrass — YUKON: on 
steep slope found on lower and mid-slope with Arabis, 
Arctostaphylos uva-ursi, Festuca saximontana and Poa, 
hill across Tagish Road from Crag Lake, 60°15'30"N 
134°28'53"W, B. Bennett 01-008, 19 May 2001 (DAO); 
summit of hill behind Carcross Cutoff, south-facing, 
pine forest Cowley Creek, 60°35.66'N 134°52.96'W, 
B. Bennett 99-300, 6 Aug. 1999 (DAO). 

Douglas et al. (1981) knew this rare species in the Territory 
from only one locality from adjacent to the Alaska Highway 
just east of the Atlin Road junction (Calder & Gillett 26477, 
DAO). Cody (1996) mapped three additional sites in the vicin- 
ity of Whitehorse. The first specimen cited above is the sou- 
thernmost yet found in the Territory. 


Vahlodea atropurpurea (Wahlenb.) Fries, Mountain 
Hairgrass — YUKON: Site YPN 120, 63°06'39.8"N 
133°20'15"W, G. Brunner 568-01, 6 Aug. 2001 (DAO). 
Cody (1996) knew this taxon from only six localities: 
adjacent to the North Canol Road, adjacent to the South 
Macmillan River and near Bennett Lake. Additional sites 
have since been found east of the Canol Road (Cody et al. 
1998 and 2000). The specimen cited above is from a site 
about 50 kilometers northeast of the Macmillan River site. 


CYPERACEAE 

Carex albo-nigra Mack., Two-toned Sedge — YUKON: 
Volcano Mountain, 62°55'N 137°23'W, G. Brunner 
521-01, 3 Aug. 2001 (DAO); moist scree with organic 
component on north-facing slope, Vulcan Mountain, 
Kluane National Park, 60°54.712'N 138°29.484'W, P. 


2004 


Caswell 527, 15 July 2002 (DAO); moist organic 
soil on southwest facing slope on old semi-abandoned 
road north of Donjek River bridge, 61°42.209'N 
139°48.376'W, P. Caswell 196, 25 June 2002 (DAO). 

Cody (1996) knew this rare species in the Territory from 
seven localities south of latitude 63°N, three of which were 
in Kluane National Park. The first specimen cited above is 
the northernmost yet found in the Territory. Cody et al. (2000) 
added another site south of Ross River and Cody et al. 
(2001) another site in the extreme southeast. 


Carex athrostachya Olney — YUKON: moist soil by 
lake, Five Mile Lake Campground, 5 km N of Mayo, 
63°39'11.1"N 135°53'14.2"W, Cody & Cody 37641, 
1 Aug. 2001 (DAO). 

Douglas et al. (1981) considered this species rare in the 
Territory on the basis of a specimen collected by R. T. Porsild 
on the shores of a small bog pond in the vicinity of Mayo, 
63°37'N 135°55'W, on 12 May 1967 (Porsild 1975). The 
specimen cited above: from nearby is only the second yet 
found in the Territory. 


Carex atrosquama Mack. — YUKON: steep gravelly 
slope adjacent to the Alaska Highway 3 km east of 
Snag Junction, 62°12'42.3"N 140°41'33.1"W, Cody 
& Cody 37861, 20 July 2002 (DAO); cleared gravel 
by highway, Top of the World Highway Km 92, 
64°6'6"N 140°46'47"W, Cody & Cody 38024, 26 July 
2002 (DAO). 

The first specimen cited above is an extension of the known 
range in the Territory about 190 kilometers northwest from the 
south end of Kluane Lake (Cody 1996) and the second spec- 
imen is an extension of the known range of about 100 kilo- 
meters west of the Dempster Highway. 


Carex capillaris L. ssp. chlorostachys (Steven) Love 
et al. — YUKON: Klondike River near mouth of Little 
Klondike, 64°02'00"N 137°41'30"W, G. Brunner 
440, 3 July 2001 (DAO). 

The specimen cited above is the first known to Cody (1996) 
between latitudes 64°N and 66°N east of the Dempster High- 
way. It is an extension of about 60 kilometers northeast of a 
site southwest of the south end of the Dempster Highway. 


Carex crawfordii Fern. — YUKON: sandy lake shore, 
Kusawa Lake, 60°34'59.2"N 136°08'26.5"W, W. J. 
Cody 37718, 13 July 2002 (DAO); meadow adjacent 
to RCMP foundation, Fort Selkirk, 62°46'34.4"N 
137°23'35.9"W, Cody & Kennedy 37749A, 16 July 
2002 (DAO). 

This species was considered rare in the Territory by Doug- 
las et al. (1981). The specimen from south of Whitehorse 
cited above is an extension of the known range in the Terri- 
tory of about 250 kilometers south of a site mapped by Cody 
(1996) from adjacent to the Pelly River. The second specimen 
is from about 85 kilometers west of a site adjacent to the 
Pelly River. 


Carex eleusinoides Turcz. — YUKON: moist organic 
soil with pumice component, down old road west of 
Alaska Highway north of Donjek River bridge, 
61°42.209'N 139°48.376'W, P. Caswell 205, 2 June 
2002 (DAO); gravelly silt near water line, Klondike 
River, 64°O1'N 137°51'W, G. Brunner 546, 8 July 


Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON VI 


565 


2001 (DAO); approximately 2 miles downstream 
from O’Brian Creek on Klondike River, 64°01'46"N 
38°01'00"W, G. Brunner 547, 15 July 2001 (DAO). 

This species was considered rare in the Territory by Doug- 
las et al. (1981). The first specimen cited above is an extension 
of the range known to Cody (1996) of about 50 kilometers 
northwest of the north end of Kluane Lake. The second and 
third specimens are the first known between latitudes 64°N 
and 66°N east of the Dempster Highway and are an extension 
of about 45 kilometers northeast of a site south of the south 
end of the Dempster Highway. 


Carex foenea Willd. — YUKON: Halfway between Order- 
ly House and Robert Luke Cabin, 62°46'34.4"N 
137°23'35.9"W, Cody & Kennedy 37773, 17 July 
2002 (DAO). 

Douglas et al. (1981) considered this species rare in the 
Territory. Cody (1996) knew it only from three sites, two of 
which were in the vicinity of Pelly Crossing. The specimen 
cited above came from the same area. Cody et al. (1998) re- 
ported an additional site in the vicinity of Beaver Ridge/ 
Larsen Creek. 


Carex limosa L. — YUKON: Sphagnum bog, 64°02'N 
137°41'W, G. Brunner 441, 3 July 2001 (DAO). 

The specimen cited above is only the second known 
between latitudes 64°N and 66°N east of the Dempster High- 
way (Cody 1996). 


Carex loliacea L. — Yukon: damp silty ground, forest 
Picea mariana, Poa glauca, Alnus incana, Salix sp., 
north side of Klondike River near Parker Creek, 
64°OI'N 137°51'W, G. Brunner 592, 9 July 2001 
(DAO); spring alongside road, gravelly, Big Gold 
Creek/60 Mile River Valley, 64°01'N 140°42'W, G. 
Brunner 116, 23 July 1991 (DAO). 

Cody (1996) considered this species uncommon in the 
Territory although mapped north to the Porcupine River. 
The specimens cited above are about equally distant east 
and west of Dawson City. 


Carex obtusata Lilj., Blunt Sedge — YUKON: British 
Mountains, 69°26'N 140°07'W, A. Martell 625, 12 July 
1979 (DAO); steep south-facing bluffs along Klondike 
River, | mile north of 64°02'00"N 137°41'30"W, G. 
Brunner 432, 2 June 2001 (DAO). 

The first specimen cited above is the northernmost yet 
found in the Territory (Cody 1996). It is an extension of the 
known range of about 80 kilometers northwest from sites 
adjacent to the Babbage River. The second specimen cited 
above is an extension of the known range in the Territory of 
about 75 kilometers southeast of a site at Sheep Mountain 
adjacent to the Dempster Highway (Kojima and Brooke 
1985). 


Carex pachystachya Cham. — YUKON: open meadow, 
abandoned beaver pond, Deadman Creek, S. Withers 
SWO1-117, 16 July 2002 (B. Bennett Herbarium, photo 
DAO) (determined by A. A. Reznicek). 

This species was not included in The Rare Vascular 
Plants of the Yukon (Douglas et al. 1981) because it was 
widespread in Canada. Cody (1996) knew it from only two 
localities in the extreme southern Yukon. The specimen 
cited above is from a site about 25 kilometers south south- 
east of Whitehorse. 


566 


Carex pellita Muhl. ex Willd. — YUKON: silty river 
bank with Carex utriculata, C. aquatilis, and Juncus 
alpinoarticulatus, Selwyn, Yukon River, 62°48'06"N 
138°15'29"W, B. Bennett 02-293, 9 Aug. 2002 (DAO); 
silty mud at edge of river at high water line with Juncus 
alpinoarticulatus, Brittania Creek, Yukon River, 
62°52'28"N 138°42'52"W, B. Bennett 02-671, 10 
Aug. 2002 (DAO); silty mud at edge of river at high 
water line with Juncus alpinoarticulatus, Kirkman 
Creek, Yukon’ River, (62°59'21;N«.139°23'07" W)) B. 
Bennett 02-675, 10 Aug. 2002 (DAO); river bank in 
area frequently scoured by river ice, confluence of 12 
Mile River, Yukon River, 64°15'11"N 139°43'18"W, 
B. Bennett 02-806, 23 Aug. 2002 (DAO); deeply 
rooted, edge of river in silty sand bank, Yukon River, 
64°33'28"N 140°38'24"W, B. Bennett 02-807, 24 
Aug. 2002 (DAO). 

Cody et al. (2000) reported this plant as a rare species in 
the Territory based on a specimen from a site below Rink 
Rapids adjacent to the Yukon River. An additional site was 
reported from Ear Lake, Whitehorse (Cody et al. 2002). The 
five sites reported above extend the known range in the Terri- 
tory about 275 kilometers northwest of the Rink Rapids. 
Bruce Bennett has found that Carex pellita is a common 
species on the Yukon River between Pink Rapids and White 
River, less common but can be found downstream and even- 
tually being replaced by C. saxatilis. 


Carex petasata Dewey — YUKON: east slope adjacent 
to Alaska Highway ca. 3 km W of Enger Creek, 
62°19'12.05"N 140°49'19.9"W, Cody & Cody 37511, 
25 July 2001 (DAO); open, abandoned mine road 
surface, Viceroy Mine — Big Rock Zone, 64°02'N 
138°17.9"W, S. Withers SWO1-057, 3 July 2002 (DAO). 
The first specimen cited above is an extension of the known 
range in the Territory (Cody 1996) of about 215 kilometers 
west of a site west of Carmacks and 245 kilometers northwest 
of a site adjacent to Haines Junction. The second specimen 
cited above is at the northern limit in the Territory about 50 
kilometers east of a site in the vicinity of Dawson City. 


Carex praegracilis Boott — YUKON: meadow adjacent 
to RCMP foundation, Fort Selkirk, 62°46'34.4"N 
137°23'35.9"W, Cody & Kennedy 37770B, 17 July 
2002 (DAO); sandy lake shore, Kusawa Lake, 
60°34'59.2"N 135°08'26.5"W, W. J. Cody 37717, 13 
July 2002 (DAO). 

Douglas et al. (1981) knew this rare species in the Terri- 
tory from only three localities: Mackintosh, Whitehorse and 
Carcross. Cody (1996) knew it from 11 sites in the Territory, 
most of which were south of latitude 61°15'N. The second 


specimen cited above is a slight extension of range south of 
Whitehorse. 


Carex rossii R. Br., Ross’ Sedge — YUKON: decom- 
posed bedrock, South Fork bluffs, Klondike River, 
64°O1'N 138°9'W, G. Brunner 446-01, 14 July 2001 
(DAO); steep south-facing bluffs along Klondike River 
1 mile north of 64°02'N 137°41'30"W, G. Brunner 
433, 2 June 2001 (DAO). 

The specimens cited above are only the third and fourth 
collections known from north of latitude 64°N (Cody 1996) 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


which are from about 80 kilometers east of a site adjacent to 
Dawson. To the south, the nearest known site is adjacent to 
the northern South Canol Road about 300 kilometers to the 
southeast. 


Carex stylosa C. A. Mey. — Cody et al. (2000) reported 
this species which is rare in the Yukon Territory from 
the Arctic Coast. 

The specimen on which it was based (Hoefs & Smitts 
93-37) was misidentified and the range extension should be 
deleted. There was also an error in the citation where the 
latitude should have been 69°23'N not 60°29'N 


Carex supina Wahl. ssp. spaniocarpa (Steud.) Hultén 
— YUKON: Steep south-facing bluffs along Klondike 
River | mile north of 64°02'N 137°41'30"W, G. Brun- 
ner 434, 2 June 2001 (DAO). 

The specimen cited above is the first known in the Territory 
north of 64°N east of the Dempster Highway. The nearest 
known site is about 75 kilometers to the northwest cited by 
Kojima & Brooke (1985). 


Carex sychnocephala Carey — YUKON: damp, fine 
gravel with organic component, at base of highway 
embankment, Alaska Highway just north of Copper 
Joe Creek, 61°19.814'N 138°56.805'W, P. Caswell 
570, 21 July 2002 (DAO). 

Douglas et al. (1981) considered this species rare in the 
Territory. The specimen cited above is an extension of the 
range known to Cody (1996) of about 175 kilometers north- 
west by west of a site west of Whitehorse. 


Carex viridula Michx. — YUKON: on moist shoreline 
of Nares Lake south of Carcross, 60°9'20"N 
134°40'1L0"W, W. J. Cody 38318, 1 Aug. 2001 (DAO). 

This species was considered rare in the Territory by Doug- 
las et al. (1981). The specimen cited above is an extension 
of the known range to Cody (1996) of about 80 kilometers 
south of Whitehorse. 


Eleocharis uniglumis (Link) Schult. — YUKON: open 
wet area in old road, adjacent to the Alaska Highway, 
61°42.209'N 139°48.376'W, P. Caswell 204, 25 June 
2002 (DAO); in wet moss by lake shore, Fish Lake, 
60°46'3.3"N 135°03'13"W, W. J. Cody 38246, 2 Aug. 
2002 (DAO). 

The species was considered rare in the Territory by Douglas 
et al. (1981). The first specimen cited above is an extension 
of the known range to Cody (1996) of about 150 kilometers 
northwest of a site northwest of Haines Junction and the 
second specimen is from a site just south of Whitehorse. 


Eriophorum brachyantherum Trautv., Short-anthered 
Cotton-grass — YUKON: Vuntut National Park, vicinity 
of Snowdrift Camp, 68°21.4'N 139°13.1'W, P. Cas- 
well PPC-2000-Y-070, 21 June 2000 (DAO). 

The specimen cited above is the northernmost yet found 
in the Territory (Cody 1996). It is an extension of about 100 
kilometers north from sites adjacent to the Porcupine River. 


Kobresia simpliciuscula (Wahlenb.) Mack., Simple 
Kobresia — YUKON: along road track through Picea 
glauca forest, Minto RV Campground, 62°35'N 
136°51'W, Cody & Cody 37678, 6 Aug. 2001 (DAO). 


2004 


The nearest site in the Territory (Cody 1996) to the speci- 
men cited above is about 130 kilometers to the southeast 
adjacent to the Klondike Highway. 


Scirpus rollandii Fern. — YUKON: moist organic soil, 
side road west of Alaska Highway, north of Donjek 
River bridge, 61°42.209'N 139°48.376'W, P. Caswell 
202, 25 June 2002 (DAO). 

Cody (1996) knew this species in the southwest of the 
Territory where he considered it rare. The specimen cited 
above is an extension of the known range of about 100 kilo- 
meters northwest of southern Kluane Lake. 


JUNCACEAE 

Juncus bufonius L. s.1., Toad Rush — YUKON: mud 
bar, Dezadeash River below Haines Junction, Kluane 
National Park, 60°45.545'N 137°34.470'W, P. Cas- 
well 791, 16 Aug. 2002 (DAO). 

The specimen cited above is new to Kluane National Park 
and is a range extension of about 290 kilometers to the south 
of a site mapped by Cody (1996) and a site east of Johnson’s 
Crossing reported by Cody et al. (2001). 


Juncus filiformis L. — YUKON: in muck by small lake 
adjacent to the Alaska Highway, 60°51'01"N 
135°46'0.5"W, Cody & Cody 38185, 1 Aug. 2002 
(DAO). 

The specimen cited above which is from a site about 75 
kilometers west of Whitehorse is the westernmost yet found 
adjacent to the Alaska Highway. 


Luzula arcuata (Wahlenb.) Sw. ssp. unalaschkensis 
(Buch) Hultén — gravel beside road and steep slope, Al- 
aska Highway Km 1918, 62°16'26.1"N 140°45'6.7"W, 
Cody & Cody 37851, 20 July 2002 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory (Cody 1996) of about 215 kilometers 
northwest of a site northwest of Haines Junction. 


LILIACEAE 
Allium schoenoprasum L. ssp. sibiricum (L.) Celak, 
Wild Onion — YUKON: Herschel Island, 69°35'N 
139°05'W, C. E. Kennedy s.n., 1 July 1985 (Environ- 
ment Yukon Herbarium, photo DAO). 

The nearest site known to Cody (1996) was adjacent to 
the Firth River at 69°13'N 139°35'W, about 30 kilometers 
southwest of the Herschel Island site. 


Maianthemum canadense Desf. ssp. interius (Fern.) 
A. & D. Love, Wild Lily-of-the- Valley — YUKON: com- 
mon in understory in the riparian white spruce poplar 
zone, Contact Creek, 60°00'00"N 127°43'44"W, B. 
Bennett 01-025, 25 June 2001 (DAO). 

Cody (1994, 1996) suggested that this taxon should be 
looked for in the Territory and Cody et al. (1998, 2000) re- 
ported it from the extreme southeast. The specimen cited 
above is an extension of the known range of about 80 kilo- 
meters southwest from a site adjacent to the Beaver River 
hotspring. 


Tofieldia coccinea Richards., Northern False Asphodel 
— YUKON: south aspect-xeric-graminoid/forb/lichen 
meadow, Volcano Mountain, 62°55'N 137°23'W, G. 
Brunner 521a-01, 3 Aug. 2001 (Environment Yukon, 
photo DAO). 


Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON YI 


567 


The specimen cited above is from the first known site 
between latitudes 62°N and 64°N. The nearest site to the north 
is about 175 kilometers to a site adjacent to the Dempster 
Highway and to the southwest about 190 kilometers adjacent 
to the Alaska Highway. 


IRIDACEAE 

Tris setosa Pall. ssp. interior (Anders.) Hultén, Wild 
Iris — YUKON: at high water level, confluence of 12 Mile 
River and Yukon River, 64°15'11"N 139°43'18"W, B. 
Bennett 02-743, 23 Aug. 2002 (DAO). 

Cody (1996) knew this rare species in the Territory from 
only three locations, two from near the western end of the 
Alaska Highway and one from adjacent to the Takhini River 
west of Whitehorse. The specimen cited above is an extension 
of the known range of about 225 kilometers to the north of the 
sites near the Alaska border. Only four plants were observed. 


ORCHIDACEAE 
Spiranthes romanzoffiana Cham. & Schlecht., Hooded 
Ladies’-tresses — YUKON: tussocks near bog, east of 
Haines Highway, 60°08.570'N 136°58.469'W, P. Cas- 
well 453, 13 July 2002 (DAO). 

The specimen cited above is an extension of the known 


range in the Territory of about 120 kilometers southwest of 
Whitehorse. 


SALICACEAE 
Salix alaxensis (Anderss.) Cov. ssp. longistylis (Rydb.) 
Hultén — YUKON: beside recreation centre in middle 
of town, Beaver Creek, 62°22'56"N 140°52'41"W, B. 
Bennett 01-102, 12 June 2001 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory (Cody 1996) of about 140 kilometers 
northwest of a site near the north end of Kluane Lake. 


Salix glauca L. var. acutifolia (Hook.) C. Schneider — 
YUKON: Mt. Merrill, 60°06'N 124°45'W, G. Brunner, 
1994 (DAO). 

The specimen cited above is about 90 kilometers east of 
the easternmost site of Salix glauca sl. mapped by Cody 
(1996). 


Salix pseudomyrsinites (Anderss.) Ball ex Hultén (S. 
novae-angliae Anderss.) — YUKON: along Bennett 
Lake just above high water mark, Carcross Dunes, 
60°10'30"N 134°43'26"W, B. Bennett & G. Argus 99- 
548, 29 Aug. 1999 (DAO) (determined by G. Argus). 

The nearest site of this species known to Cody (1996) is in 
the vicinity of Whitehorse, about 60 kilometers to the north- 
west. 


Salix rotundifolia Trauty. ssp. dodgeana (Rydb.) Argus 
— YUKON: Dryas alaskensis tundra, Mount Casca 
border Monument 97, 65°21'25"N 141°00'00"W, B. 
Bennett & M. B. Cook 02-519, 27 Jane 2002 (DAO) 
(determined by G. Argus). 

The specimen cited above from adjacent to the Alaska 
border is from a site about 70 kilometers northwest of the 
nearest site known to Cody (1996). 


URTICACEAE 

Urtica dioica L. ssp. gracilis (Ait.) Selander, Sting- 
ing Nettle — YUKON: dwelling mound, Forty Mile 
Historic Site at the confluence of Forty Mile and 


568 


Yukon rivers, 64°25'N 140°30'W, C. E. Kennedy 44, 
July 2000 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory (Cody 1996) of about 50 kilometers 
northwest of the vicinity of Dawson City. 


POLYGONACEAE 

Polygonum lapathifolium L., Willow Weed, Pale Smart- 
wood — YUKON: wetland Carex fen, uncommon silty 
mud beside beaver pond, 60°07'00"N 124°15'2"W, B. 
Bennett 98-609, 16 June 1998 (DAO). 

This species was considered rare in the Territory by Doug- 
las et al. (1981). The specimen cited above is an extension 
of the known range in the Territory of about 700 kilometers 
southeast of sites in the vicinity of Carmacks. 


Rheum rhabarbarum L. (R. rhaponticum L.) Rhubarb 
— YUKON: adjacent to garage near machine shop, Fort 
Selkirk, 62°46'34.4"N 137°23'35.9"W, Cody & Ken- 
nedy 37799, 17 July 2002 (DAO). 

Cody et al. (2001) and (2002) reported Rhubarb growing 
wild at Silver City and Destruction Bay. The specimen cited 
above is the third known cited where it was growing wild. 


Rumex crispus L., Curled Dock — YUKON: moist shore 
of small lake between Ross River and Campbell 
Highway, 61°58'7"N 132°38'34"W, Cody & Cody 
37961, 24 July 2002 (DAO). 

The specimen cited above is the sixth site in the Territory 
for this introduced species. Cody (1996) knew it from the 
vicinity of Dawson City and it has since been reported from 
just west of Whitehorse, Haines Junction and adjacent to the 
Alaska Highway near the Alaska border (Cody et al. 2002). 


CHENOPODIACEAE 

Atriplex alaskensis S. Watson, (A. patula sensu Cody 
(1996), A. patula var. alaskensis (S. Wats.) Welsh), 
Alaskan Orache (Figure 3) — YUKON: alkaline flat, 
Takhini Salt Flats, 60°51.2'N 135°43.23'W, Bennett & 
Parker 98-431, 2 Sept. 1998 (B. Bennett Herbarium, 
photo DAO) (determined by P. W. Ball); alkaline flats, 
between the salt flat and the highway in the area of the 
fence, Takhini Salt Flats, 60°51'23"N 135°42'55"W, 
B. Bennett 01-168, 23 Aug. 2001 (B. Bennett Herb- 
arium, photo DAO); occasional in wet alkaline margin 
of a brackish slough, Mile 484.5 Alaska Highway west 
of Whitehorse, R. L. Taylor 4094, 14 July 1959 (DAO); 
common around clay margins of dried up alkaline 
pond, east of Haines Junction at Mile 985 Alaska 
Highway, 60°48'N 136°45'W, Calder & Kukkonen 
25252, 14 Aug. 1960 (DAO); common around clayey 
margin of alkaline lake, between Minto and Pelly 
Crossing on Dawson-Whitehorse road, approximately 
62°46'N 136°36'W, Calder & Kukkonen 28095, 10 
Aug. 1960 (DAO). 

Cody (1996) considered the Calder & Cody specimens as 
introductions from Europe (A. patula). Atriplex alaskensis 
should now be added to the Yukon Flora and the list of rare 
plants of the Territory (Douglas et al. 1981). The Taylor spe- 
cimen cited above was the basis for the drawing published 
in the Yukon Flora as A. patula. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Atriplex subspicata (Nutt.) Rydberg — YUKON: grassy 
area in open alkaline salt ponds, Mayo Road, Dilla- 
bough’s Grazing Lease, 60°55'54"N 135°10'14"W, B. 
Bennett 02-649, 19 Aug. 2002 (DAO). 

Cody (1996) knew this species from a single locality adja- 
cent to a saline pond just west of Whitehorse. The specimen 
cited above from just north of Whitehorse is only the second 
known from the Territory. 


Chenopodium capitatum (L.) Asch., Strawberry-blite 
— YUKON: disturbed gravelly soil, roadstop, Atlin Road 
just north of BC border, 60°00'09"N 133°47'42.1"W, 
Cody & Cody 37546, 28 July 2001 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory (Cody 1996) of about 60 kilometers 
to the southwest from a site in the vicinity of Johnson’s 
Crossing. 


Corispermum ochotense Ignatov var. alaskanum 
Mosyakin (C. hyssopifolium sensu Cody 1996) — 
YUKON: river bar in loose aluvial sand between Salix 
exigua and river with Tanacetum bipinnatum, Poten- 
tilla anserina and Aster falcatus, Yukon River at con- 
fluence of Nester Creek, 64°38'22"N 140°52'52"W, 
Bennett & Mulder 02-665, 25 Aug. 2002 (DAO); river 
bar in loose alluvial sand between Salix exigua and 
river with Tanacetum bipinnatum, Potentilla anserina 
and Aster falcatus, Yukon River, 64°22'19"N 
140°26'49"W, Bennett & Mulder 02-740, 24 Aug. 
2002 (DAO). 

The specimens cited above are the first yet found in the 
Territory. 


PORTULACACEAE 

Claytonia megarrhiza (A. Gray) Parry, Alpine Spring- 
beauty (Figure 3) — YUKON: alpine tundra, three plants 
seen at base of a long scree slope near a very small 
stream, northwest side of Wade Mountain, Kluane 
National Park, 61°18'N 139°33'W, P. Caswell 236, 
28 June 2002 (DAO). 

This species, which is new to the Flora of the Yukon Terri- 
tory (Cody 1996), should be added to the list of rare plants 
of the Territory (Douglas et al. 1981). Porsild and Cody 
(1980) knew it from three sites in the Mackenzie Mountains 
in the Continental Northwest Territories and McJannet et al. 
(1995) knew it from four sites in that area in the Rare 
Vascular Plants in the Northwest Territories. To the south 
Douglas et al. (2002) mapped three sites in the extreme 
southeast of British Columbia where it was considered rare. 


Montia fontana L., Blinks — YUKON: aquatic, growing 
in the shallows (10-30 cm) in a backwater slough, 
Lewes Marsh, 60°34'N 134°35'W, B. Bennett OO- 
1091, 25 June 2000 (DAO) (determined by C. Parker 
and W. J. Cody). 

Cody (1996) knew this rare plant in the Territory only from 
a single locality on the Arctic Coast where it was collected 
by Erling Porsild in 1934 (CAN, photo DAO). In addition a 
second specimen was collected by Cody on Herschel Island 
in 1999 (Cody et al. 2001). This is a circumpolar, low-arctic 
species, which was considered rare in the Territory by 
Douglas et al. (1981). To the east of the location cited above 
it is known from the southern Mackenzie Mountains in the 


2004 


FiGure 3. Claytonia megarrhiza, Alpine Springbeauty (drawn 
by Lynne Bartosch). 


former District of Mackenzie, and to the west in essentially 
coastal regions of Alaska and British Columbia. 


CARYOPHYLLACEAE 
Cerastium arvense L., Field Chickweed — YUKON: wet 
roadside ditch, Top of the World Highway Km 100, 
64°05'53"N 140°54'43"W, Cody & Cody 38011, 26 
July 2002 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory (Cody 1996) of about 100 kilometers 
west of the vicinity of Dawson City. 


Minuartia dawsonensis (Britt.) House — YUKON: steep 
slope by Alaska Highway, 8 km west of White River, 
62°3'15.6"N 140°38'20.5"W, Cody & Cody 37880, 
20 July 2002. 

The specimen cited above is an extension of the known 
range in the Territory (Cody 1996) of about 125 kilometers 
northwest of a site adjacent to Kluane Lake. 


Minuartia elegans (Cham. & Schlecht.) Schischk. — 
YUKON: Dryas alaskensis heath, 30° southwest-facing 
large talus dolomite boulders, summit of mountain 
on Canadian side of Mount Casca, 65°21'37"N 
140°59'34"W, B. Bennett & M. B. Cook 02-543, 27 
June 2002 (B. Bennett Herbarium, photo DAO). 

The specimen cited above from adjacent to the Alaska bor- 
der is an extension of the known range of about 75 kilometers 
northwest of the nearest site in the Ogilvie Mountains. 


Minuartia macrocarpa (Pursh) Ostenf. — YUKON: al- 
pine tundra, spotty occurrence in raised rocky mounds, 
mountain between Kusawa and JoJo Lake, 60°35'47"N 
136°15'19"W, B. Bennett 97-681, 19 Sept. 1997 (DAO). 
The specimen cited above is an extension of the known 
range in the southwest of the Territory of about 80 kilo- 
meters southeast of a site northwest of Haines Junction. 


Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON VI 


569 


Minuartia yukonensis Hultén — YUKON: coarse gravel 
and small rocks of a stream’s outwash, Upper Joe 
Creek, Kluane National Park, 61°12.861'N 139°03.863'W, 
P. Caswell 585, 24 July 2002 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory (Cody 1996) of about 225 kilometers 
west of a site north of Whitehorse. 


Silene involucrata (Cham. & Schlecht.) Bocquet ssp. 
tenella (Tolm.) Bocquet — YUKON: meadow between 
Stone House and river near Fort Selkirk sign, Fort 
Selkirk, 62°46'34.4"N 137°23'35.9"W, Cody & Ken- 
nedy 37805, 17 July 2002 (DAO). 

The specimen cited above is from a site between sites in 
the vicinity of Mayo and south of latitude 62°. 


Silene williamsii Britton — YUKON: talus lower slope 
of southwest-facing bluff, Yukon River, Minto Bluff, 
62°37'06"N 136°57'41"W, B. Bennett 02-678, 9 Aug. 
2002 (B. Bennett Herbarium, photo DAO); common 
on base of slope across from Brittania Creek growing 
with Artemisia frigida and Calamagrostis purpuras- 
cens, Yukon River, 62°52'28"N 138°42'56"W, Bennett 
02-443, 10 Aug. 2002 (B. Bennett Herbarium, photo 
DAO). 

This species was considered rare in the Territory by 
Douglas et al. (1981) on the basis of 3 collections from south 
and southwest of Dawson City. 


Silene vulgaris (Moench) Garcke, Bladder Campion 
— YUKON: gravelly soil in old field with grass and 
sweet clover, west of Mayo airport, 63°37'01.4"N 
135°58'20.9"W, Cody & Cody 37665, 4 Aug. 2001 
(DAO). 

The specimen cited above of this introduced species in 
the Territory (Cody 1996) was previously known only from 
an area just east of Dawson. 


Stellaria umbellata Turcz. — YUKON: wet scree cushion 
in zinc moss, Wade Mountain, Kluane National Park, 
61°18.450'N 139°30.921'W, P. Caswell 354, 7 July 
2002 (DAO). 

Douglas et al. (1981) considered this species as rare in the 
Territory. Cody (1996) knew it from only three widely sepa- 
rated sites. The specimen cited above is from a site about 60 
kilometers northwest of a previously known site in Kluane 
National Park. 


NYMPHAEACEAE 
Nuphar polysepalum Engelm., Yellow Pond-lily — 
YUKON: shallow water along shore of lake, Tatchun 
Lake Campground, 62°17'52.8"N 136°08'11.4"W, 
Cody & Cody 37626, 30 July 2001 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory (Cody 1996) of about 120 kilometers 
to the north, south of Mayo. 


RANUNCULACEAE 

Pulsatilla ludoviciana (Nutt.) Heller, Prairie-crocus — 
YUKON: steep grassy slope on ridge northeast of Wood- 
burn Creek, Tintina Trench, Ddhaw Ghro, 63°08'N 
136°05'W, C. E. Kennedy 4, 27 July 2001 (DAO). 


570 


The specimen cited above is an extension of the known 
range in the Territory (Cody 1996) of about 100 kilometers 
east northeast of a site north of the junction of the Pelly and 
Yukon rivers. 


Ranunculus aquatilis L. var. subrigidus (W. B. Drew) 
Breitung — YUKON: near South Fork intake, Klondike 
River on edge of beaver dam in old ditch, 64°0'30"N 
138°12'00"W, G. Brunner 549, 22 July 2002 (DAO). 

The nearest site known to Cody (1996) of this uncommon 
variety was about 60 kilometers to the north adjacent to the 
Dempster Highway. 


Ranunculus occidentalis Nutt. var. brevistylis Greene, 
Western Buttercup — YUKON: alpine tundra, valley 
above treeline, Macmillan Pass, 63°15'N 130°02'W, 
J. Basinger s.n., June 1982 (DAO). 

Douglas et al. (1981) considered this species rare in the 
Territory. The specimen cited above is the northernmost yet 
found in the Territory and is an extension of the known 
range of about 250 kilometers northeast of a site adjacent to 
the northern South Canol Road. 


PAPAVERACEAE 

Papaver croceum Ledeb. — YUKON: by garbage along 
trail in Picea glauca, Pinus, Populus tremuloides wood- 
land, Whitehorse, near Hidden Lake, 60°35'00.5"N 
135°51'00.2"W, Cody & Cody 37678A, 9 Aug. 2001 
(DAO); in gravel beside car parking spot, 11 Chalet 
Crescent, Whitehorse, 60°55'54.7"N 135°10'26.5"W, 
W. J. Cody 37711, 12 July 2002 (DAO) (determined 
by H. Solstad). 

This species is a garden escape which has not previously 
been recorded from the Whitehorse area. In addition Heidi 
Solstad revised three collections previously determined as 
Papaver nudicaule (Dawson, Calder & Billard 3121 (DAO), 
Halfway Lakes area 15 miles north of Mayo, Calder & Gil- 
lett 4176 (DAO) and Tower Hill, Tagish, R. Rosie 798 (DAO)) 
to P. croceum. P. nudicaule ssp. nudicaule should be replaced 
by P. croceum in the flora (Cody 1996, 2000). 


BRASSICACEAE (CRUCIFERAE) 

Alyssum obovatum (C. A. Meyer) Turez. (A. ameri- 
canum Greene) — YUKON: 45° talus southwest-facing 
slope, Yukon River, | km east of Alaska/Yukon border, 
64°41'05"N 140°57'30"W, Bennett & Mulder 02-666, 
25 Aug. 2002 (B. Bennett Herbarium, photo DAO) 
(determined by G. A. Mulligan). 

Douglas et al. (1981) considered this species too wide- 
spread to include it in The Rare Vascular Plants of the Yukon. 
Cody (1996) knew it from only eleven sites north of latitude 
62°N. The specimen cited above is a slight extension of the 
known range in the Territory about 50 kilometers to the 
northwest of another site adjacent to the Yukon River. 


Arabis caucasica Willd., Wall Rock-Cress (Figure 4) 
— YUKON: garden escape, Alsek Road, Hayes Property, 
10 km NW of Haines Junction, 64°47'N 137°41'W, 
B. Bennett 01-041, 12 June 2001 (DAO) (determined 
by G. A. Mulligan). Submitted to B. Bennett by 
Carolyn Hayes. 


This garden escape has not previously been observed in 
the Yukon Territory. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


re a 


Ze 
AL 


Figure 4. Arabis caucasica (drawn by Lee Mennell). 


Arabis caucasica can be separated from A. codyi as 
follows: 


A. Garden escape; cauline and caudex leaves 
with similar dentate to subdentate margins, 
flowering stems spreading to ascending; 
petals!/6:Simim) longi ee ee ae A. caucasica 


B. Native species; cauline and caudex leaves not 
similarly dentate to subdentate; flowering stems erect; 
petals more than 10 mm long............. A. codyi 


Arabis holboellii Hornem. var. retrofracta (Graham) 
Rydb. — YUKON: gravel beside road and steep slope, 
Alaska Highway Km 1918, 62°16'26.1"N 140°45'6.7"W, 
Cody & Cody 37855, 20 July 2002 (DAO). 

The nearest site adjacent to the Alaska Highway known 
to Cody (1996) is about 150 kilometers to the southeast in the 
vicinity of Kluane Lake. There is, however, a site adjacent 
to a river about 60 kilometers to the east. 


Arabis holboellii Hornem. var. secunda (Howell) Jep- 
son — YUKON: adjacent to narrow road through woods 
leading down to Nares Lake southeast of Carcross, 
60°9'20"N_ 134°40'10"W, Cody & Cody 37934, 23 
July 2002 (DAO) (determined G. A. Mulligan). 

Cody (1996) knew this taxon from only four localities in 
the Territory, west of longitude 135°. Cody (2001) added new 
sites in the vicinity of Whitehorse and Wolf Lake to the east. 
The specimen cited above is the southernmost yet found in 
the Territory. 


, 
{ 
' 


2004 


Armoracia rusticana (Lam.) Gaert., Mess. & Scherb. — 
YUKON: Hultén (1941-50) reported this species “A specimen 
of this plant was collected at Stewart R. July 1898 by Anders- 
son (S). It must have been cultivated or have escaped from 
cultivation". Hultén (1968) mapped the worldwide distribution 
of this species with a single dot in the Territory and Welsh 
(1974) sub Rorippa armorocia (L.) A. S. Hitche. stated “Cul- 
tivated for the root; in southern Alaska and Yukon, persisting: 
introduced from Europe". Fortunately this species was over- 
looked by Cody when preparing the Flora of the Yukon 
Territory (1996); since the Anderson specimen was recently 
borrowed from the Swedish Museum of Natural History, 
Stockholm and was revised by Gerald A. Mulligan to 
Rorippa barbareifolia (DC.) Kitagawa, an amphi-Berigian, 
nonarctic species which extends across Alaska to central 
Yukon Territory, the Porcupine River valley, the central Rich- 
ardson Mountains and the Fort McPherson area south of the 
Mackenzie Delta. 


Brassica rapa L., Bird rape — YUKON: roadside gravel, 
Alaska Highway, near Kluane Wilderness Camp Km 
1791, B. Bennett 01-095, 24 July 2001 (DAO). 

The specimen cited above of this introduced species is an 
extension of the known range in the Territory of about 175 
kilometers northwest of a site adjacent to the Haines High- 
way. It may have resulted from roadside seeding. 


Camelina sativa (L.) Cranz, Falseflax (Figure 5) — 
YUKON: disturbed site, Whitehorse Shipyards, 
60°43'34"N 135°03'12"W, B. Bennett 01-150, 28 Aug. 


RNA 


tS a Men 
AP, 
SS Yee 
“has ls 
yy 
AY 
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FiGure 5. Camelina sativa, Falseflax (drawn by Lee Mennell). 


Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON VI 


S74 


2001 (B. Bennett Herbarium, photo DAO) (determined 
by G. A. Mulligan). 

The specimen cited above is new to the Flora of the Yukon 
Territory (Cody 1996). It is a Eurasian introduction which is 
known in Canada from New Brunswick to British Columbia 
and north into southern District of Mackenzie. 

The genus Camelina can be separated from the genus 
Alyssum as follows: 


A. — Styles usually 2-3 mm long; silicles oval-elliptic to 
egg-shaped, usually at least 5 mm long; seeds numer- 
ous; petals often shallowly bilobed; annual Camelina 


B. Styles not over 1 mm long; silicles oval 
in outline, less than 5 mm long; petals 
notbilobed:perennialy ys ee eae ere Alyssum 


Capsella bursa-pastoris (L.) Medic., Shepherd’s-purse 
— YUKON: level gravel area, Campbell Highway Km 
380, 62°02'34.2"N 132°52'02.2"W, Cody & Cody 
37591, 30 July 2001 (DAO); growing beside Artemi- 
sia tilesii by narrow road through woodland, Pelly 
Farm Road, 62°51'22.2"N 137°00'34"W, Cody & 
Kennedy 37821B, 17 July 2002 (DAO); gravel, Pick- 
handle Lake, 63°02'22.2"N 138°23'15.8"W, Cody & 
Cody 37834, 20 July 2002 (DAO). 

The first specimen cited above of this introduced species 
is from a site between two sites adjacent to the Campbell 
Highway (Cody 1996) in the vicinity of Carmacks. One of 
sites is 175 kilometers to the west and the other is about 185 
kilometers to the southeast. The second and third specimens 
are from sites between Keno and Carmacks (Cody 1996). 


Cardamine bellidifolia L., Alpine Bittercress — YUKON: 
Herschel Island, 69°35'N 139°O5S'W, C. E. Kennedy 
s.n., 20 July 1985 (DAO). 

The nearest sites known to Cody (1996) were from the 
north coast about 30 kilometers to the west and near the 
Firth River about 30 kilometers to the southwest. 


Descurainia incisa (Engelm. ex A. Gray) Britton var. 
incisa, Tansy Mustard — YUKON: gravel roadside, rare, 
Dempster Highway Km 98, 64°42'N 138°24'46"W, 
Cody & Cody 38056, 27 July 2002 (DAO) (deter- 
mined by G. A. Mulligan). 

Cody et al. (2001) reported this species as new to the 
Territory and Cody et al. (2002) added additional sites adja- 
cent to the Alaska Highway north of latitude 62°N. The 
specimen cited above is from the first known site adjacent 
to the Dempster Highway. 


Draba aurea M. Vahl, Golden Draba — YUKON: Site 
KP 110, 62°15\09-6°N) 137°41:03) W. (Gs Brunner 
591-01, 8 Aug 2001 (Environment Yukon Herbariam, 
photo DAO) (determined by G. A. Mulligan). 

The specimen cited above is the northernmost yet found 
in the Territory (Cody 1996). It is an extension of the known 
range of about 50 kilometers from a site south of latitude 
62°N. 

Draba corymbosa R. Br. — YUKON: Herschel Island, 
69°35'N 139°05'W, C. E. Kennedy s.n., 10 Aug. 1985 


(Environment Yukon Herbarium, photo DAO) (deter- 
mined by G. A. Mulligan). 


S72 


The nearest site known to Cody (1996) was from near the 
Firth River about 30 kilometers to the southwest. 


Draba crassifolia Graham — YUKON: sandy moist soil in 
eroded area, ridge south of Donjek Glacier, 61°08.277'N 
139°31.170'W, P. Caswell 403, 8 July 2002 (DAO) 
(determined by G. A. Mulligan). 

The specimen cited above is an extension of the known 
range in the Territory (Cody 1996) northwest of sites in 
Kluane National Park. 


Draba glabella Pursh — YUKON: steep south-facing 
bluffs along Klondike River, 1 mile north of 
64°02'00"N_ 137°41'30"W, G. Brunner 435, 2 June 
2001 (DAO) (determined by G. A. Mulligan). 

The specimen cited above is from a site about 70 kilo- 
meters east of a site in the vicinity of Dawson City (Cody 
1996). 


Draba nemorosa L., Wood Whitlow-grass — YUKON: 
disturbed farm area, Black Sheep airline base, Mayo, 
63°35.26'N 135°51.83'W, B. Bennett 00-1193, 2 July 
2000 (DAO) (determined by G. A. Mulligan). 

The specimen cited above is an extension of the known 
range in the Territory (Cody 1996) of about 110 kilometers 
northeast of a site west of longitude 137°W and north of the 
Pelly River. 


Draba ogilviensis Hultén — YUKON: summit of small 
peak south of Donjek Glacier, Kluane National Park, 
61°08.318'N 139°31.132'W, P. Caswell 392, 8 July 
2002 (DAO) (determined by G. A. Mulligan). 

Douglas et al. (1981) considered this species rare in the 
Territory. The specimen cited above is the westernmost yet 
known in Kluane National Park. 


Draba palanderiana Kjellm. — YUKON: dry alpine 
Dryas heath with Dryas alaskensis, Silene acaulis, 
Synthyris borealis, Podistera macounii and lichens, 
summit of mountain, calcareous dolomite talus, 
65°21.37'N 140°59.34'W, B. Bennett 02-523, 27 June 
2002 (DAO). 

The specimen cited above from adjacent to the Alaska 
border is an extension of the known range in the Territory of 


about 60 kilometers to the north of a site mapped by Cody 
(1996). 


Draba porsildii G. A. Mulligan — YUKON: wet scree, 
Wade Mountain, Kluane National Park, 61°18'N 
139°33'W, P. Caswell 240, 2 June 2002 (DAO) (deter- 
mined by G. A. Mulligan). 

Douglas et al. (1981) considered this species rare in the 
Territory on the basis of three specimens from Kluane Nation- 
al Park and one from the South Canol Road southwest of Ross 
River. Cody (2003) cited a specimen from west of the three 
sites in Kluane National Park. The specimen cited above is 
from an additional site in the Park about 70 kilometers north 
of the last site. 


Draba stenoloba Ledeb. — YUKON: Donjek Valley, 
61°25.689'N 139°53.074'W, R. Maraj s.n., 29 June 
2002 (DAO) (determined by G. A. Mulligan). 

The specimen cited above is an extension of the known 
range in the Territory (Cody 1996) of about 90 kilometers 
northwest of a site in Kluane National Park. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Lepidium densiflorum Schrad. var. densiflorum, Com- 
mon Pepper-grass — YUKON: roadstop in disturbed 
gravelly soil, Atlin Road just north of BC border, 
60°00'09"N_ 133°47'42.1"W, Cody & Cody 37543, 
37551, 28 July 2001 (DAO) (determined by G. A. 
Mulligan). 

The specimens cited above are an extension of the known 
range in the Territory (Cody 1996) of about 125 kilometers 
southeast of a site just west of Whitehorse. 


Lepidium densiflorum Schrad. var. macrocarpum G. 
A. Mulligan — YUKON: adjacent to Catholic Church, 
Fort Selkirk, 62°46'34.4"N 137°23'35.9"W, Cody & 
Kennedy 37808A, 17 July 2002 (DAO) (determined by 
G. A. Mulligan). 

Cody et al. (2001) reported this variety, which is probably 
introduced, new to the Territory from the vicinities of Daw- 
son, Ross River and Pelly Crossing. The specimen cited 
above is an extension of the known range in the Territory of 
about 50 kilometers to the west of Pelly Crossing. 


Parrya arctica R. Br. — Two specimens, one from Pauline 
Cove, Herschel Island (Cody 36030) and the other from the 
Bonnet Plume River (Loewen 99-25-93) were recently revised 
to P. nudicaulis by G. A. Mulligan. These two species can 
be separated as follows: 


A. Stamens linear, 1.5 to 
IEPfoyistVool) KON reteieicn acc bane jaa P. nudicaulis 


B. Stamens oblong, less than 
1h Oimmvlon es kane een OT ate ee P. arctica 


In addition, Cody (1994) reported P. arctica from Herschel 
Island on the basis of a specimen collected by P. F. Cooper 
(433), 9 May 1979 (CAN) as new to the Territory and sug- 
gested that it should be added to the list of rare plants (Doug- 
las et al. 1981). This specimen has now been revised to Thlas- 
piarcticum Porsild by G. A. Mulligan and should be deleted 
from the flora of Herschel Island and the Yukon Territory. 


Rorippa barbareifolia (DC.) Kitagawa — YUKON: Pelly 
Crossing, 62°51'09"N 136°56'05.2"W, Cody & Ken- 
nedy 37831, 17 July 2002 (DAO) (determined by G. 
A. Mulligan). 

This species was considered rare in the Territory by Doug- 
las et al. (1981). The specimen cited above is the southern- 
most yet found adjacent to the Alaska Highway. 


Sinapis arvensis L., Charlock — YUKON: in a well-tend- 
ed flower bed, Haines Junction, P. Caswell 311, 5 July 
2002 (DAO) (determined by G. A. Mulligan). 

The specimen cited above which was the only plant ob- 
served in Haines Junction of this introduced species was from 
only the third site known to Cody (1996) in the Territory. 


Subularia aquatica L. ssp. americana Mulligan & 
Calder, Awlwort — YUKON: shallows of large pond east 
of Alaska Highway north of Sulphur Lake, 61°00.435'N 
138°12.729'W, P. Caswell 772, 9 Aug. 2002 (DAO) 
(determined by G. A. Mulligan). 

This species was considered rare in the Territory by Doug- 
las et al. (1981). The specimen cited above is an extension 
of the known range of about 175 kilometers east of a site in 
the vicinity of Whitehorse (Cody 1996). 


2004 


Thlaspi arvense L., Penny Cress — in black muck just 
inside gate, flats 7 miles east of Tagish Bridge, 
60°19'S"N 134°10'17"W, Cody & Cody 37958, 23 
July 2002 (DAO) (determined by G. A. Mulligan). 

The specimen cited above is an extension of the known 
range of this introduced species of about 80 kilometers south- 
east of the vicinity of Whitehorse. 


SAXIFRAGACEAE 
Ribes oxyacanthoides L. ssp. oxyacanthoides — YUKON: 
meadow adjacent to RCMP foundation, Fort Selkirk, 
62°46'34.4"N 137°23'35.9"W, Cody & Kennedy 37757, 
16 July 2002 (DAO). 

Although quite frequent in the Territory south of latitude 
62°N, Cody (1996) knew this shrub from only two sites to 
the north. 


Saxifraga caespitosa L.— YUKON: 40° slope talus dolo- 
mite boulders, Mount Casca border Monument 97, 
65°21'25"N 141°00'00"W, B. Bennett & M. Cook 02- 
SISA, 27 June 2002 (DAO). 

Cody (1996) mapped this species in the Territory from 
three distinct areas (between latitudes 60° and 62°, between 
latitudes 64° and 66°, and north of latutude 68°. The specimen 
cited above from adjacent to the Yukon-Alaska border is about 
70 kilometers from the sites east and southeast. 


Saxifraga eschscholtzii Sternb. — YUKON: dry alpine 
heath at summit of mountain on Canadian side of 
Mount Casca, 65°21'37"N 140°59'34"W, B. Bennett 
& M. Cook 02-512, 27 June 2002 (DAO). 

Cody (1996) knew this species only from the British 
Mountains in the extreme northwest of the Territory. The 
specimen cited above is an extension of the known range in 
the Territory of about 460 kilometers to the south. 


Saxifraga foliolosa R. Br. — YUKON: damp sand of 
small sandy depression in sedge tundra, ridge south 
of Donjek Glacier, Kluane National Park, 61°07.383'N 
139°31.308'W, P. Caswell 416, 8 July 2002 (DAO). 

Douglas et al. (1981) considered this species which was 
known to Cody (1996) from north of latitude 67°N and a 
single site on the west side of Haines Highway southeast of 
Haines Junction as rare in the Territory. Cody et al. (2000) 
reported another site in the south from about 80 kilometers 
northeast of the site adjacent to the Haines Highway. The 
specimen cited above is an extension of the range in the south 
of about 135 kilometers northwest of the site near the Haines 
Highway. 


Saxifraga nelsoniana D. Don ssp. pacifica (Hultén) — 
YUKON: wet organic soil with some fine scree at edge 
of stream, Tabletop Mountain, Kluane National Park, 
61°15.231'N 139°11.115'W, P. Caswell 285, 29 June 
2002 (DAO). 

Douglas et al. (1981) considered this subspecies rare in 
the Territory. The specimen cited above is an extension of the 
known range in the Park of about 25 kilometers west of a site 
west of Kluane Lake. 


Saxifraga rufopilosa (Hultén) Porsild — YUKON: moist 
places in 30° southwest facing large talus dolomite 
boulders, summit of mountain on Canadian side of 
Mount Casca, 65°21'37"N 140°59'34"W, B. Bennett 
& M. B. Cook 02-528, 27 June 2002 (DAO). 


Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON VI 


SME: 


The specimen cited above is from a site about 35 kilometers 
northwest of the northernmost site south of latitude 66°N 
known to Cody (1996). 


Saxifraga tricuspidata Rottb., Prickly Saxifrage — 
YUKON: steep grassy slope on ridge northeast of Wood- 
burn Creek, Tintina Trench, Ddhaw, 63°08'N 136°05'W, 
C. E. Kennedy 7, 27 July 2001 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory (Cody 1996) of about 100 kilometers 
east northeast of a site at about longitude 137°W, north of the 
Pelly River. 


ROSACEAE 
Amelanchier alnifolia (Nutt.) Nutt., Saskatoon — 
YUKON: southwest-facing slope across from Brittania 
Creek, Yukon River, 62°52'28"N 138°42'56"W, B. 
Bennett 02-653, 10 Aug. 2002 (DAO). 

Cody (1996) knew only three sites of this species north of 
the site reported above: Dawson City, Mayo, and adjacent 
to the Stewart River. 


Chamaerhodos erecta (L.) Bge. ssp. nuttallii (Picker- 
ing ex Rydb.) Hultén — YUKON: common at base of 
southwest-facing slope, across from Brittania Creek, 
Yukon River, 62°52'28"N 138°42'56"W, B. Bennett 
02-640, 10 Aug. 2002 (DAO). 

Cody (1996) knew this species in the Territory north to 
near latitude 64°. The specimen cited above is an extension 
of the known range in the Territory of about 80 kilometers 
west of a site adjacent to the Pelly River. 


Dryas hookeriana Juz. — YUKON: Ogilvie Mountains, 
Tombstone Range, Yakamaw Creek, watershed east of 
Angelcomb Peak, 64°36'N 138°14'W, Cody 36808, 
36845, 20-22 July 1999 (DAO); Dryas alaskensis 
tundra, Mount Casca border Monument, 65°21'25"N 
141°00'00"W, B. Bennett & M. B. Cook 02-518, 27 
June 2002 (B. Bennett Herbarium, photo DAO). 

The specimens cited above are the first yet known to Cody 
(1996) from west of the Dempster Highway between latitudes 
64°N and 66°N. 


Geum aleppicum Jacq. ssp. strictum (Ait.) Clausen, 
Yellow Avens — YUKON: Top of the World Golf Course, 
64°3'38"N 139°26'13"W, Cody & Cody 37998, 25 
July 2002 (DAO). 

Douglas et al. (1981) considered this taxon rare in the Ter- 
ritory. The specimen cited above is an extension of the known 
range in the Territory of about 200 kilometers west of a site 
west of Keno City (Cody et al. 2001) and north of sites adja- 
cent to the Alaska Highway near the Alaska border. 


Geum rossii (R. Br.) Scr. — YUKON: moist places at sum- 
mit of the Canadian side of Mount Casca, 65°21'37"N 
140°59'34"W, B. Bennett 02-524, 27 June 2002 (B. 
Bennett Herbarium, photo DAO). 

The specimen cited above is from a site about 100 kilo- 
meters northwest of the nearest location in the Ogilvie Moun- 
tains where it has been collected frequently. 


Potentilla arguta Pursh ssp. convallaria (Rydb.) Keck, 
Tall Cinquefoil — YUKON: dry rocky area, South Fork 
intake, Klondike River, 64°0'30"N 138°12'00"W, G. 
Brunner 551, 22 July 2002 (DAO). 


574 


The specimen cited above is only the second collected 
from north of latitude 64°N in the Territory (Cody 1996). It is 
from a site about 60 kilometers east of the vicinity of Dawson. 


Rosa woodsii Lindl., Western Rose — YUKON: Artemi- 
sia frigida slope, east side of lake, Whitehorse, Hid- 
den Lakes, 60°41'30"N 135°02'45"W, B. Bennett O1- 
118, 11 July 2001 (B. Bennett Herbarium, photo 
DAO); grassy alkaline slope north of highway on slope, 
Alaska Highway Km 1588, | km east of Cracker Creek, 
60°48'20"N 136°48'00"W, B. Bennett 02-735, 11 Sept. 
2002 (DAO); common at base of slope, Yukon River, 
Brittania Creek, 62°52'28"N 138°42'56"W, B. Bennett 
02-639, 10 Aug. 2002 (B. Bennett Herbarium, photo 
DAO). 

This is a rare species in the Territory (Douglas et al. 1981). 
The first specimen cited above is only the fourth known in 
the Territory (Cody 1996). The second is an extension of the 
known range in the Territory (Cody 1996) of about 50 kilo- 
meters west of a site adjacent to the Alaska Highway. The 
third specimen is an extension of about 125 kilometers north- 
west of a site reported by Cody et al. (2003) from the vic- 
inity of Carmacks. 


FABACEAE (LEGUMINOSAE) 

Lupinus nootkatensis Donn — YUKON: clearing bet- 

ween road and poplar forest, west side of Haines High- 

way, north of cut-off to Dalton Post, 60°09.405'N 

136°58.572'W, P. Caswell 440, 13 July 2002 (DAO). 
The specimen cited above is only the third known from 

southeastern Kluane National Park and the Yukon Territory. 


Oxytropis arctica R. Br., Arctic Oxytrope — YUKON: at 
base of talus slope, Lower Kathleen Lake, along south 
shore ca. 25 km SE of Haines Junction, 60°33'N 
137°16'W, G. W. & G. G. Douglas 8391, 25 June 
1975 (Kluane National Park, photo DAO); in alpine 
fell field, Dezadeash River Valley ca. 16 km WSW of 
Haines Junction, G.W. & G.G. Douglas 8405, 30 June 
1975 (Kluane National Park Herbarium, photo DAO). 

Cody (1996) suggested that this species should be looked 
for in the mountains of northern Yukon Territory because it 
is known in the Canadian Arctic and northern Alaska. Douglas 
et al. (1999) included the Territory in the distribution of O. 
arctica presumably on the basis of the specimens cited above 
and also reported the only collection known in British Colum- 
bia from Mile 416 Alaska Highway which is about 700 kilo- 
meters east southeast of the Yukon sites. It should be added 
to the list of rare species in the Territory. 


Oxytropis campestris (L.) DC. ssp. varians (Rydb.) 
Cody, Field Locoweed — YUKON: steep grassy slope 
on ridge northeast of Woodburn Creek, Tintina Trench, 
Ddhaw Ghro, 63°08'N 136°05'W, C. E. Kennedy 6, 
27 July 2001 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory (Cody 1996) of about 110 kilometers 
northeast of a site adjacent to the Yukon River. 


Trifolium pratense L., Red Clover — YUKON: road- 
side, junction of Campbell Highway and Frenchman 
Lake Road, 62°4'30"N 135°30'55"W, Cody & Cody 
37985, 24 July 2002 (DAO). 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


The nearest site of this introduced species in the Territory 
(Cody 1996) is from the vicinity of Whitehorse, about 150 
kilometers to the south. 


Vicia americana Muhl., American Vetch — YUKON: bor- 
der of grassland and Populus, Macoobs Park, Mayo, 
63°35'42.5"N 135°56'57.1"W, Cody & Cody 37668, 
5 Aug. 2001 (DAO); west end of parking lot by visitor 
kiosk, Rancheria, 60°05'16"N 130°36'10"W, B. Ben- 
nett 01-059, 26 July 2001 (DAO). 

Douglas et al. (1981) considered this species rare in the 
Territory on the basis of a single collection from the Larsen 
Creek hotsprings in the extreme southeast (Scotter & Cody 
1979). Additional collections were added from this area by 
Cody et al. (1998) and from the vicinity of Watson Lake (Cody 
et al. 2000). The first specimen cited above is an extension 
of the known range in the Territory of about 550 kilometers 
to the northwest of Watson Lake. The second collection is an 
extension of the known distribution of about 120 kilometers 
west of Watson Lake but was probably introduced at that site. 


CALLITRICHACEAE 
Callitriche anceps Fern. — YUKON: on mud substrate at 
edge of water, old beaver pond east of Haines Highway 
about 10 kilometers south of Kathleen Lake cutoff, 
60°30.838'N 137°04.970'W, P. Caswell 787, 11 Aug. 
2002 (DAO). 

This species was considered rare in the Territory (Douglas 
et al. 1981). The specimen cited above is from a site about 
30 kilometers northwest of a site in Kluane National Park. 


VIOLACEAE 

Viola renifolia Gray var. brainerdii (Greene) Fern., 
Kidney-leaved Violet - YUKON: humus covered with 
old poplar leaves, poplar and white spruce wood flat, 
Wade Lakes Road, between Kluane National Park and 
Haines Highway, 60°07.119'N 137°05.369'W, P. Cas- 
well 020, 6 June 2002 (DAO). 

Cody (1996) knew this species as far north as the vicinity 
of Dawson City. The specimen cited above is an extension 
of the known range in the Territory of about 150 kilometers 
southwest of the vicinity of Whitehorse. 


HALORAGACEAE 

Myriophyllum verticillatum L. — YUKON: in water be- 
side dock, Nunatuk Camp, 62°9'49"N 135°48'4"W, 
Cody & Cody 37989, 24 July 2002 (DAO). 

Douglas et al. (1981) considered this species rare in the 
Territory. The specimen cited above is from a site about half 
way between the Pelly River and the north end of Lake Le- 
barge. 


APIACEAE 

Cicuta maculata L. var. angustifolia Hook., Spotted 
Water-hemlock — YUKON: lush herbaceous meadow on 
riverbank, Kirkman Creek, Yukon River, 62°59'21"N 
139°23'07"W, B. Bennett 02-284, 10 Aug. 2002 
(DAO). 

Douglas et al. (1981) considered this taxon rare in the 
Territory. The specimen cited above is an extension of the 
known range of about 175 kilometers to the northwest of 
Carcross (Cody 1996). 


Podistera macounii (Coult. & Rose) Mathias & Const. 
— YUKON: Dryas alaskensis tundra, Mount Casca bor- 


2004 


der Monument 97, 65°21'25"N 141°00'00"W, Bennett 
& Cook 02-520, 27 June 2002 (DAO). 

This species was considered rare in the Territory by Doug- 
las et al. (1981). Cody (1996) knew it only from the Richard- 


_ son Mountains area in the northeast. Cody et al. (1998) ex- 


tended the known range in the Territory south into the Daw- 
son range at latitude 62°26'N. The specimen cited above is 
an extension of the known range in the Territory of about 
275 kilometers to the west from the southernmost site in the 
Richardson Mountains. 


PYROLACEAE 

Pyrola minor L. — YUKON: adjacent to the Catholic 

Church, Fort Selkirk, 62°46'34.4"N_ 137°23'35.9"W, 

Cody & Kennedy 37815, 17 July 2002 (DAO). 
Although quite frequent in the Territory east and north of 

the Klondike Highway, Cody (1996) also knew this species 


_ from the vicinity of southern Kluane Lake, to the west. 


_ ERICACEAE 


Cassiope tetragona (L.) D. Don ssp. saximontana 
(Small) Porsild — YUKON: dry alpine Dryas heath, 
summit of Mount Casca, 65°21'37"N 140°59'34"W, 
Bennett & Cook 02-522, 27 June 2002 (DAO). 


The specimen cited above is the northernmost yet found in 


_ the Territory. The nearest site to the above north of Chapman 


Lake in the Ogilvie Mountains is about 125 kilometers to the 
southeast (Cody et al. 2001). 


| Harrimanella stellariana (Pallas) Coville — YUKON: 
Morley River watershed, Englishman Range, 8 km SW 


of Morris Lake, northwest facing slope growing in 
bedrock controlled draw, 60°23'40"N 131°47'50"W, 
J. Meikle 02-126, 20 June 2002 (DAO). 

The specimen cited above is a rarity in the Territory (Doug- 
las et al. 1981). It is a slight range extension north of the 
eastern site mapped by Cody (1996). 


Phyllodoce glanduliflora (Hook). Cov., Yellow Moun- 
tain-heather — YUKON: subalpine slope by creek, Tank 
Creek, 60°05'30"N 134°41.9'W, S. Withers SWO1-114, 
15 July 2001 (DAO). 

Cody (1996) knew this species in the Territory only from 
sites east of longitude 130°W and west of longitude 137°W. 
The specimen cited above is about 125 kilometers east of a 
site in Kluane National Park. 


Vaccinium membranaceum Dougl., Tall Blueberry — 
YUKON: moist subalpine meadow, Kotaneelee Range, 
60°14'31L"N 124°7'19"W, B. Bennett 95-149, 20 June 
1998 (B. Bennett Herbarium, photo DAO); subalpine 
forest, Beavercrow Ridge, 60°12.865'N 124°35.79'W, 
B. Bennett 98-310, (B. Bennett Herbarium, photo 
DAO). 

Cody (1996) suggested that this species should be looked 
for in the southeastern part of the Territory. Cody et al. (1998) 
reported four collections from Mount Merrill and Gusty 
Lakes. The two specimens cited above are from the extreme 
southeast. 


PRIMULACEAE 
Douglasia alaskana (Cov. & Standl.) Kelso — YUKON: 
eroded alpine slope, Table Mountain, Kluane National 


Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON VI 


575 


Park, 61°15'.231'N 139°11.115'W, P. Caswell 282, 
29 June 2002 (DAO). 

Douglas et al. (1981) considered this species rare in the 
Territory where it was only known in Kluane National Park. 
The specimen cited above extends the known range in the 
Park about 75 kilometers to the northwest. 


Douglasia arctica Hook. — YUKON: Cody et al. (2000) 
reported this species from a mountain between Kusawa 
and JoJo Lake, which was a considerable southward 
range extension. 

The specimen upon which it was reported was unfortu- 
nately misidentified and the range extension should be deleted. 


Primula eximia Greene (P. tschuktschorum Kjellm. 
ssp. cairnesiana A. E. Porsild) — YUKON: Dempster 
Highway about Km 96, at about altitude 4000 ft., S. 
Frisch, spring 2001 (DAO). 

Douglas et al. (1981) considered this species rare in the 
Territory on the basis of collections from the northern Rich- 
ardson Mountains and a site adjacent to the Alaska border at 
about 63°50'N latitude. The specimen cited above is from a 
site about 150 kilometers northeast of the Alaska border site 
where there was “a whole field of them". 


Trientalis europaea L. s.1., Starflower — YUKON: damp 
mossy Picea glauca, Alnus incana, Dawson area, 
64°02'00"N 137°41'30"W, G. Brunner 439, 30 June 
2001 (DAO). 

The specimen cited above from about 70 kilometers east of 
Dawson City, is only the second collection in the Territory 
from north of latitude 64°N (Cody 1996). 


GENTIANACEAE 

Gentianella tenella (Rottb.) Boerner — YUKON: alpine 
barrens on long ridge, Atlas Mountain, Kluane Nation- 
al Park, 61°14.772'N 139°19.254'W, D. Normandeau 
s.n., P- Caswell 328A, 6 July 2002 (DAO); patch of al- 
pine tundra, mountain opposite Donjek Glacier, Kluane 
National Park, 61°11.859'N 139°23.372'W, P. Cas- 
well 619, 26 July 2002 (DAO). 

This species was considered rare in the Territory by Doug- 
las et al. (1981). Cody (1996) knew it from only three sites, 
two about 40 kilometers west of the specimens cited above 
and one just north of latitude 62°N. 


POLEMONIACEAE 
Phlox hoodii Richards., Moss Phlox — YUKON: dry 
rocky slope, open habitat, Donjek River watershed near 
Kluane Glacier, M. Hoefs 02-693, 4 July 2002 (DAO). 
The specimen cited above from west of the Alaska High- 
way is the fifth and southernmost in that area. This species 
was not included in the Rare Vascular Plants of the Yukon 
(Douglas et al. 1981) because of its widespread distribution. 


Polemonium acutiflorum Willd. forma lacteum Lepage 
— YUKON: rare in lush subalpine meadow at 4800 ft.. 
mountain 4 miles west of Upper Hyland Lake, 62°03'N 
128°59'W, Calder & Kukkonen 27889, 3 Aug. 1960 
(DAO) (determined by B. Boivin 1970). 

Lepage (1950) described this white-flowered form on the 
basis of a specimen collected at Anchorage, Alaska. Cody 
(1996) unfortunately did not mention this white form 
occurring in the Yukon Flora. 


576 


Polemonium boreale Adams forma albiflorum Cody, 
— YUKON: fox den, sandy/grass area, Herschel Island, 
site 203, 69°30'N 139°15'W, C. E. Kennedy 262, 16 
July 1985 (DAO) (PARATYPE); [NORTHWEST TERRI- 
TORIES: MACKENZIE: on a sandy portion of a gravel bar, 
Horton River, N.W.T., 69°42'N 126°56'W, G. W. 
Scotter 101016b, 6 July 1995 (DAO) (HOLOTYPE) 
(Cody et al. 2003). 

Cody (1996) stated that the corolla of this species was 
blue to violet or rarely white. The white form had not been 
described at that time. 


HYDROPHYLLACEAE 
Phacelia mollis Macb., Macbryde’s Phacelia — YUKON: 
scattered in dry, open rubble on south-facing slopes 
near base of bluff, Moosehide Hills, Bluff on N side 
of Yukon R., 25 km downstream from Dawson, 
64°15'41"N 139°36'13"W, C. Roland 93-27, 24 May 
1993 (AKA, photo DAO). 

The specimen cited above is an extension of about 50 kilo- 
meters to the southwest from the nearest site in the Ogilvie 
Mountains known to Cody (1996). 


BORAGINACEAE 

Eritrichium aretioides (Cham. & Schlecht.) DC. — 
YUKON: dry alpine Dryas heath, Canada/US Boundary 
Monument 97, 65°21'25"N 141°00'00"W, B. Bennett 
02-019, 27 June 2002 (DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 40 kilometers northwest of the 
nearest site known to Cody (1996) between latitudes 64° 
and 66°N. 


Eritrichium chamissonis DC. — YUKON: vicinity of 
Snowdrift Camp, Vuntut National Park, 68°21.4'N 
139°13.1'W, P. Caswell PPC-Y-076, 19 June 2000 (B. 
Bennett Herbarium, photo DAO). 

Cody (1996) knew this rare species in the Territory from 
only two localities in the British Mountains. The specimen 
cited above is from a site about 50 kilometers south of a site 
on Mt. Sedgwick. 


Mertensia paniculata (Britt.) G. Don var. alaskana 
(Britt.) Williams — YUKON: adjacent to Joe Robert’s 
Cabin, Fort Selkirk, 62°46'34.4"N 137°23'35.9"W, 
Cody & Kennedy 37796, 17 July 2002 (DAO). 

Cody (1996) knew this variety only from south of latitude 
61°30'N and north of latitude 67°30'N. 


Myosotis scorpioides L., Forget-me-not — YUKON: 
flower bed at Kluane Park Inn, Haines Junction, 
60°44'N 137°31'W, P. Caswell PPC-2000-Y-426, 28 
Aug. 2000 (B. Bennett Herbarium, photo DAO). 

This introduced species was previously known in the Ter- 
ritory only from the vicinity of Whitehorse (Cody 1996). 


LAMIACEAE (LABIATAE) 

Galeopsis tetrahit L. ssp. bifida (Boenn.) Fries, Hemp- 
nettle — YUKON: Forty Mile Historic Site at the conflu- 
ence of the Forty Mile and Yukon rivers, 64°25'N 
140°32'W, C. E. Kennedy 2, 6 July 2000 (DAO); under 
large Picea glauca in squirrel midden, Judas Creek, 
60°23.4'N 134°07.4'W, S. Withers SWO1-178, 30 Aug. 
2001 (B. Bennett Herbarium, photo DAO). 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


This introduced species was previously known in the Ter- 
ritory from only three localities: North Canol Road (Cody 
1994, 1996), Km 9 Dempster Highway (Cody et al. 1998) 
and Whitehorse (Cody et al. 2001). The first specimen cited 
above is the northernmost yet found in the Territory. 


SCROPHULARIACEAE 
Euphrasia subarctica Raup — YUKON: stony soil and 
moss in cleared area below rest stop, Klondike High- 
way Km 628, 63°50'18.1"N 137°59'20.5"W, Cody & 
Cody 37660, 4 Aug. 2001 (DAO). 

The specimen cited above is the northernmost yet found in 
the Territory (Cody 1996). It is from a site about 110 kilo- 
meters northwest/west of Mayo. 


Mimulus guttatus DC., Yellow Monkeyflower — 
YUKON: small seep near the road, Km 4.5 Atlin Road, 
60°17'30"N 133°50'3"W, S. Withers SWO1-131, 24 
July 2001 (DAO). 

Douglas et al. (1981) considered this species rare in the 
Territory. The specimen cited above is from a site about 80 
kilometers southeast of Whitehorse. 


Pedicularis oederi Vahl., Oeder’s Lousewort — YUKON: 
Herschel Island, 69°35'N 139°05'W, C. E. Kennedy 
s.n., 30 July 1985 (Yukon Renewable Government 
Herbarium, photo DAO). 

This species, which is frequent in the British Mountains 
to the south is now known to the north on Herschel Island. 


Rhinanathus minor L. ssp. borealis (Sterneck) A. Love 
— Yukon: gravel parking area at foot of trail to Rock 
Glacier, Haines Highway, 60°27'18"N 137°03'37.6"W, 
Cody & Cody 37894, 21 July 2002 (DAO). 

This taxon is known in the Territory only as far north as 
about latitude 61°30'N. The specimen cited above is the first 
record from the Haines Highway. 


OROBANCHACEAE 

Orobanche fasciculata Nutt. — YUKON: 45° silty sand 
south-facing slope with Artemisia frigida and Cala- 
magrostis purpurascens, Dutch Bluff, Yukon River, 
61°55.72'N 135°03.99'W, B. Bennett 02-290, 7 Aug. 
2002 (DAO). 

This species was considered rare in the Territory by Doug- 
las et al. (1981). Cody et al. (2003) reported a new site in 
Kluane National Park adjacent to the Alaska Highway and 
southern Kluane Lake. The specimen cited above is from a 
site about 50 kilometers southeast of Carmacks. 


LENTIBULARIACEAE 
Pinguicula villosa L. — YUKON: in Sphagnum tus- 
sock, bog east of Haines Highway, 60°08.421'N 
136°58.499'W, P. Caswell 446, 13 July 2002 (DAO). 
The specimen cited above is new to southwestern Yukon. 
It is an extension of the known range in the Territory of about 
300 kilometers southwest from a site adjacent to the northern 
South Canol Road. 


Utricularia minor L., Lesser Bladderwort — YUKON: 
shallow water of lake between Ross River and Camp- 
bell Highway, 61°58'16"N 132°38'06"W, Cody & 
Cody 37967, 24 July 2002 (DAO). 

This is a rare species in the Territory that Cody (1996) knew 
from only four sites between latitudes 60°N and 64°30'N. 


2004 


Cody et al. (1998, 2000 and 2003) reported new collections 
from the southeast in the vicinity of La Biche River and in 
the far north between Trout Lake and the Babbage River. 
The specimen cited above is from about 150 kilometers 
north of a site near the south end of the South Canol Road. 


PLANTAGINACEAE 
Plantago canescens Adams — YUKON: base of cliffs, 
talus, south-facing slope, Mount Carmacks, 64°20'08"N 
140°12'25"W, B. Bennett 02-820, 24 Aug. 2002 
(DAO). 

The specimen cited above is an extension of the known 
range in the Territory of about 200 kilometers northwest of 
a site adjacent to the Klondike Highway (Cody 1996). 


_ Plantago maritima L., Seaside Plaintain — YUKON: dry 


sandy gravel slope north of gravel pit south of Alaska 
Highway, 60°49'58"N 135°45'49"W, Cody & Cody 


— 38204, 1 Aug. 2002 (DAO). 


This species was considered rare in the Yukon Territory by 
Douglas et al. (1981). Cody (1996) knew it from only two 
localities adjacent to Kluane Lake in the extreme southwest. 


_ The specimen cited above is from a site about 100 kilometers 


to the east. 


_ CAPRIFOLIACEAE 
_ Sambucus racemosa L. ssp. pubens (Michx.) House, 
_ Red Elderberry — YUKON: roadside and lakeshore, 


Wade Lakes, 60°05.0'N 137°19.6'W, R. Maraj s.n., 
14 Sept. 2002 (DAO). 

Douglas et al. (1981) (as var. arborescens) knew this taxon 
from a single locality in southern Kluane National Park. The 
specimen cited above extends the known range about 80 kilo- 


_ meters to the east between the park and Haines Highway. 


VALERIANACEAE 
Valeriana sitchensis Bong., Sitka Valerian — YUKON: 
Site YPN 120, 63°06'39.8"N 133°20'15"W, G. Brun- 
ner 569-01, 6 Aug. 2001 (Environment Yukon, photo 
DAO). 

The specimen cited above is from a site between the North 
Canol Road and Mayo. 


ASTERACEAE (COMPOSITAE) 

Aster alpinus L. ssp. vierhapperi Onno, Alpine Aster 
— YUKON: steep grassy slope on ridge, northeast of 
Woodburn Creek, Tintina Trench, Ddhaw Ghro, 


_ 63°08'N 136°05'W, C. E. Kennedy 2, 27 July 2001 
_ (DAO). 


The specimen cited above is an extension of the known 
range in the Territory (Cody 1996) of about 100 kilometers 
northeast of a site near the junction of the Pelly and Yukon 
rivers. 


Aster falcatus Lindl., Western Heath Aster — YUKON: 
lush herbaceous meadow riverbank, Kirkman Creek, 
Yukon River, 62°59'21"N 139°23'07"W, B. Bennett 
02-673, 10 Aug. 2002 (B. Bennett Herbarium, photo 
DAO). 

Douglas et al. (1981) considered this species rare in the 
Territory. The specimen cited above is from a site about 150 
kilometers west of a site adjacent to the Klondike Highway. 


Erigeron caespitosus Nutt. — YUKON: silty sand south- 
facing slope, Dutch Bluff, Yukon River, 61°55.72'N 


Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON VI 


yd 


135°03.99'W, B. Bennett 02-300, 7 Aug. 2002 (DAO); 
top of slope of open bank, 1 mile upstream of Big 
Salmon, on Yukon River, 61°51.41'N 134°55.01'W, 
B. Bennett 02-302, 7 Aug. 2002 (DAO); base of cliffs, 
talus, south-facing slope, Yukon River, Mount Car- 
macks, 64°20'08"N 140°12'25"W, B. Bennett & R. 
Mulder 02-816, 24 Aug. 2002 (DAO). 

The first specimen cited above is from a site about 75 kilo- 
meters southeast of a site adjacent to Carmacks and the sec- 
ond specimen is from a site about 120 kilometers southeast 
of Carmacks. The third specimen is from a site between two 
sites near the Alaska border, one to the south about 120 kilo- 


meters and the other to the north about 100 kilometers (Cody 
1996). 


Erigeron hyperboreus Greene — YUKON: middle of 
dirt road leading to Copper Joe Creek Road, Kluane 
National Park, 61°18.688'N 138°56.319'W, P. Caswell 
170, 24 June 2002 (DAO). 

This species was considered as rare in the Territory by 
Douglas et al. (1981). The specimen cited above is an exten- 
sion of about 425 kilometers south of a site northwest of 
Dawson City. 


Senecio eremophilus Richards., Dryland Ragwort (Fig- 
ure 6) — YUKON: Pine Creek Campground, 4 miles 
east of Haines Junction, Mile 1012 Alaska Highway, 
V. L. Harms 6299, 17 August 1968 (DAO). 

The specimen cited above is the first known record in the 
Yukon Territory where it was probably introduced. To the 


FiGurE 6. Senecio eremophilus, Dryland Ragwort (drawn 
by J. Looman). 


578 


west in Alaska, Hultén (1968) knew it only from the 
vicinity of Tok where he considered it also to be introduced. 
To the east Porsild and Cody (1980) knew it from the Slave 
River area north of Fort Smith where it was found in damp 
woodland meadows and adjacent to roadsides and considered 
native. To the south it is found in eastern British Columbia 
east to Manitoba and south into the United States. It can be 
separated from S. sheldonensis as follows: 


A. Stem leaves deeply incised, not 
TeducedipwardSey ss yale. coset S. eremophylus 


B. Stem leaves with merely repand-denticulate 
margins, reduced upwards ......... S. sheldonensis 


Solidago canadensis L. var. salebrosa (Piper) Jones — 
YUKON: beside test plot, 1 km N of Montague Road- 
house, Km 132 Klondike Highway, 61°49'35"N 
136°7'52"W, Cody & Cody 38073, 28 July 2002 
(DAO). 

Douglas et al. (1981) considered this species rare in the 
Territory. The specimen cited above is from about 90 kilo- 
meters southwest of a site mapped by Cody (1996). 


Sonchus arvensis L. ssp. uliginosus (Bieb.) Nyman, 
Perennial Sow-thistle — YUKON: roadside gravel, Alaska 
Highway 3 km west of Kusawa Lake turnoff below 
microwave tower, 60°46'00"N 136°05'36"W, _ B. 
Bennett 02-749, 11 Sept. 2002 (DAO). 

The specimen of this introduced species cited above is from 
a site between Whitehorse and Haines Junction. 


Tanacetum vulgare L., Common Tansey — YUKON: in 
distributed ground along Campbell Highway, Tuchitua 
River, 60°56'N 129°13'00"W, R. Rosie 1959, 16 Aug. 
1997 (DAO); single patch west of the boat launch, 
Yukon River Bridge, 60°34'20"N 134°40'W, B. Ben- 
nett 01-063, 19 Aug. 2001 (DAO). 

The specimens cited above are only the second and third 
records of this introduced species in the Territory. Cody et al. 
(1998) reported the first collection from just north of Kath- 
leen Lake Lodge on the Haines Highway. 


Acknowledgments 

We thank Donald Britton and Daniel Brunton for 
their identification of /soetes; Stephen J. Darbyshire for 
his identification of several Poaceae; Gerald Mulligan 
for his identification of Brassicaceae specimens; Heidi 
Solstad for her identification of Papaver croceum; Greg 
Brunner, Manfred Hoefs, Jennifer Line, R. Maraj, 
Rhonda Rosie and Stu Withers for their contributions; 
Margaret Cody and Douglas Cody for assisting the 
senior author in the summers of 2001 and 2002 in 
southern Yukon; artist Lee Mennell for his art work; 
Paul Catling for reviewing an earlier version of this 
manuscript, and especially Leslie Cody for the many 
hours inputting this information on her computer. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Literature Cited 

Cody, W. J. 1994. The Flora of the Yukon Territory: Addi- 
tions, Range Extensions and Comments. Canadian Field- 
Naturalist 108: 428-479. 

Cody, W. J. 1996. Flora of the Yukon Territory. National 
Research Council (NRC) Press, Ottawa, Ontario, Canada. 
643 pages. 

Cody, W. J. 2000. Flora of the Yukon Territory — Second 
Edition. National Research Council (NRC) Press, Ottawa, 
Ontario, Canada. 669 pages. 

Cody, W. J., C. E. Kennedy, and B. Bennett. 1998. New 
Records of Vascular Plants in the Yukon Territory. Cana- 
dian Field-Naturalist 112: 289-328. 

Cody, W. J., C. E. Kennedy, and B. Bennett. 2000. New 
Records of Vascular Plants in the Yukon Territory I. Cana- 
dian Field-Naturalist 114: 417-443. 

Cody, W. J., C. E. Kennedy, and B. Bennett. 2001. New 
Records of Vascular Plants in the Yukon Territory HI. 
Canadian Field-Naturalist 115: 301-322. 

Cody, W. J., C. E. Kennedy, B. Bennett, and V. Loewen. 
2002. New records of Vascular Plants in the Yukon Terri- 
tory IV. Canadian Field-Naturalist 116: 446-474. 

Cody, W. J., C. E. Kennedy, B. Bennett, and J. Staniforth. 
2003. New records of Vascular Plants in the Yukon Ter- 
ritory V. Canadian Field-Naturalist 117: 278-301. 

Cody, W. J., K. L. Reading, and J. L. Line. 2003. Additions 
and range extensions to the vascular plant flora of the Con- 
tinental Northwest Territories and Nunavut, Canada II. 
Canadian Field-Naturalist 117(3): 448-465. 

Douglas, G. W., G. W. Argus, H. L. Dickson, and D. F. 
Brunton. 1981. The rare vascular plants of the Yukon. 
Syllogeus 28: 1-96. 

Hulten, E. 1941-1950. Flora of Alaska and Yukon, 1-10. 
Lunds Universitets Arsskrift, N. F. Aud. 2. Volumes 37- 
46. 1902 pages. 

Hulten, E. 1968. Flora of Alaska and neighboring territories. 
Stanford University Press. Stanford, California. 1008 pages. 

Kojima, S., and R. C. Brooks. 1985. An annotated vascular 
flora of areas adjacent to the Dempster Highway, central 
Yukon Territory. I. Pteridophyta, Gymnospermae and 
Monocotyledonae. British Columbia Provincial Museum, 
Contributions of Natural Science 31: 1-16. 

Lepage, E. 1950. Variations mineures de quelques plantes 
du nord-est du Canada et de l’ Alaska. Le Naturaliste cana- 
dien 77 : 228-231. 

McJannet, C. L., G. W. Argus, and W. J. Cody. 1995. Rare 
vascular plants in the Northwest Territories. Syllogeus 
73: 1-104. 

Porsild, A. E. 1975. Materials for a flora of central Yukon 
Territory. National Museum of Natural Sciences. Publica- 
tions in Botany 4, Ottawa, Ontario. 77 pages. 

Porsild, A. E., and W. J. Cody. 1980. Vascular plants of Con- 
tinental Northwest Territories, Canada. National Museum 
of Natural Sciences, Ottawa, Ontario. 667 pages. 

Scotter, G. W., and W. J. Cody. 1979. Interesting vascular 
plants from southeastern Yukon Territory. Canadian Field- 
Naturalist 93: 163-170. 

Welsh, S. L. 1974. Anderson’s Flora of Alaska and Adjacent 
Parts of Canada. Brigham Young University Press, Provo, 
Utah, U.S.A. 724 pages. 


Received 11 December 2003 
Accepted 3 February 2004 


Hybridization Between a Green Turtle, Chelonia mydas, and 
Loggerhead Turtle, Caretta caretta, and the First Record of a 
Green Turtle in Atlantic Canada 


MICHAEL C. JAMES!, KATHLEEN MARTIN2, and PETER H. DuTTON? 


' Department of Biology, Dalhousie University, 1355 Oxford St., Halifax, Nova Scotia B3H 4J1 Canada (Corresponding 
author) 

> Nova Scotia Leatherback Turtle Working Group, 2070 Oxford St., Halifax, Nova Scotia B3L 2T2 Canada 

3 Southwest Fisheries Science Center, National Marine Fisheries Service, 8604 La Jolla Shores Dr., La Jolla, California 
92037 USA 


James, Michael C., Kathleen Martin, and Peter H. Dutton. 2004. Hybridization between a Green Turtle, Chelonia mydas, 
and Loggerhead Turtle, Caretta caretta, and the first record of a Green Turtle in Atlantic Canada. Canadian Field- 
Naturalist 118 (4): 579-582. 


The Green Turtle (Chelonia mydas) principally occupies tropical and subtropical waters, although juveniles are known to 
occur seasonally in temperate coastal waters. Collaboration with commercial fishers in eastern Canada yielded the most 
northerly records of this species in the northwest Atlantic. Here we report on the first confirmed record of a Green Turtle in 
eastern Canada and on the occurrence of a rare Green Turtle—Loggerhead Turtle (Caretta caretta) hybrid. Hybridization 
between the Carettini and Chelonini is extraordinary given that these groups have been genetically distinct for 50 million 


years Or more. 


Key Words: Green turtle, Chelonia mydas, hybrid, Loggerhead Turtle, Caretta caretta, Atlantic Canada. 


The Green Turtle (Chelonia mydas) has a broad 
range in the Atlantic, which includes waters off the 
continental United States. Presence in temperate waters 
of the northeastern United States is seasonal, with tur- 
tles retreating to more southerly latitudes when water 
temperatures decline (Epperly et al. 1995). During 
summer and fall, this species regularly occurs as far 
north as New York (Morreale et al. 1992); however, 
records of Green Turtles at higher latitudes of the 
United States are rare. While both Leatherback Tur- 
tles (Dermochelys coriacea) and Loggerhead Turtles 
(Caretta caretta) are commonly encountered in waters 
off Atlantic Canada (e.g., Bleakney 1965), with leath- 
erbacks occupying both nearshore and offshore waters 
(James et al. 2005) and loggerheads mainly offshore 
waters, there were no previous confirmed reports of 
the Green Turtle in this region. Here we report on 
photo-documented records of a Green Turtle and a 
Green Turtle-Loggerhead Turtle hybrid in nearshore 
waters off Nova Scotia, Canada. These exist as the 
most northerly confirmed records of Chelonia mydas 
in the northwest Atlantic. 

Notification about both turtles initially came through 
calls to a toll-free phone line established for fishing 
community members in Atlantic Canada to report sea 
turtle sightings (Martin and James 2005). A small, 
live cheloniid turtle was reported on 8 August 1999. 
The turtle was found in Chedabucto Bay, Nova Scotia 
(at 45°20'37"N, 61°15'36"W, Figure 1) and was pho- 
tographed and released shortly before the call was 


46 - (ag 


Chedabucto Bay 


450 Nova Scotia 2 4 


St Margarets 


Ite Bay 
447 ; 
| eo | 


a = A 


FiGurE 1. Capture locations of juvenile Green Turtle (solid 
circle) and juvenile Green Turtle x Loggerhead Tur- 
tle hybrid (solid triangle). 


placed. The turtle was not measured; however, curved 
carapace length (CCL) was estimated to be between 
30 and 40 cm and its mass approximately 4-5 kg. 
Species identification was confirmed as C. mydas (juve- 
nile) upon receipt of several excellent photographs 
that depicted carapacial scute configuration and scale 
patterning on the head (Figure 2). 

The second turtle, also live, was reported on 2 Octo- 
ber 2001. It was found in St. Margarets Bay, Nova Sco- 
tia (at 44°34'56"N, 64°03'06"W, Figure 1). The animal 


579 


580 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


FiGure 2. Juvenile Green Turtle (Chelonia mydas) found in Chedabucto Bay, Nova Scotia, on 8 August 1999. 


was considered unusual by the inshore commercial 
fishers who encountered it because they had never seen 
a cheloniid turtle before. Therefore, they collected it 
and brought it to shore for examination. 

One of us (MCJ) responded to the report and exam- 
ined the turtle (CCL 34 cm; mass 4.07 kg), tentatively 
identifying it as a juvenile C. mydas. The plastron of 
the turtle was cream coloured, the cutting edge of the 
lower tomium was mildly serrated, and it had a pair 
of large prefrontal scutes on its head (Figure 3), all 
features characteristic of a Green Turtle. However, 
while the number and arrangement of carapacial scutes 
was consistent with those of a Green Turtle, there 
were two claws on the anterior margin of each front 
flipper, the costal and vertebral scutes overlapped con- 
siderably, and the marginal scutes were strongly ser- 
rated (Figure 3), which suggested that the turtle was 
possibly a hybrid. 

At the time of examination, the turtle’s movements 
were sluggish and it was judged to be mildly hypother- 
mic. Small juvenile cheloniid turtles foraging in coastal 
areas of the temperate north Atlantic in the fall are 
particularly vulnerable to developing hypothermia, as 
water temperatures can rapidly drop below 20°C (Dav- 
enport 1997). This animal was recovered from an area 
where the sea surface temperature was 16.2°C. Given 
the turtle’s compromised physical condition and the 
declining water temperatures along the coast of Nova 
Scotia, the turtle was moved to the animal care facili- 


ties at Dalhousie University in Halifax, where it was 
warmed and rehydrated for approximately 60 hours 
in a freshwater bath at 24°C. It was then transported 
south by air on the evening of 4 October 2001 for 
release in Bermudian waters. 

Subsequent genetic analysis at the U.S. National 
Marine Fisheries Laboratory in La Jolla, California, 
confirmed that this turtle was a hybrid. This was deter- 
mined by sequencing a 391 bp fragment of the control 
region of mitochondrial DNA (mtDNA) extracted 
from a blood sample (Dutton 1996). The results revealed 
that this turtle had Loggerhead Turtle mtDNA, while 
phenotypically it was principally a Green Turtle. Since 
mtDNA is maternally inherited, we conclude that this 
animal was the progeny of a female Loggerhead Turtle 
and a male Green Turtle. 


Discussion 

The extent to which natural hybridization in sea tur- 
tles occurs has not been determined. However, as genet- 
ic techniques like the ones employed here are increas- 
ingly applied, hybridization may prove to be more 
common than previously thought. Molecular genetics 
have confirmed hybridization between the Loggerhead 
Turtle and Kemp’s Ridley Turtle (Lepidochelys kempii) 
(Karl et al. 1995; Barber et al. 2003), Loggerhead 
Turtle and Hawksbill Turtle (Eretmochelys imbricata) 
(Karl et al. 1995; Witzell and Schmid 2003), and Green 
Turtle and Hawksbill Turtle (Wood et al. 1983; Karl 


2004 


et al. 1995). In addition to the results presented here, 
there is only one previous report of hybridization 
between a Loggerhead Turtle and a Green Turtle (Karl 
et al. 1995). Hybridization between the Carettini and 
Chelonini is extraordinary given that these tribes have 
been genetically separated for 50 million years or more 
(Bowen et al. 1993; Dutton et al. 1996). It has been 
suggested that these may be the oldest vertebrate line- 
ages known to hybridize in nature (Karl et al. 1995). 

The reproductive status of marine turtle hybrids is 
not known; however, the identification of potential 
second-generation hybrids (Karl et al. 1995) suggests 
that at least some hybrids may be fertile. Reproductive 
viability in hybrids could have important biological 
consequences for the conservation of marine turtles 
(Karl 1996). 

Both turtles reported here were of the size class typ- 
ical of neritic foraging populations of Green Turtles 
(i.e., straight carapace length >25 cm) (Musick and 
Limpus 1997). Therefore, these animals likely arrived 
in eastern Canadian waters from coastal foraging habi- 
tat off the northeastern United States, rather than from 
pelagic areas. 

Our current understanding of juvenile Green Turtle 
distribution in the northwest Atlantic suggests that these 
turtles occupy areas where sea surface temperature is 
normally higher than that of coastal Atlantic Canada. 
In the case of the hybrid turtle, the Loggerhead Turtle 
component of its genotype may have been responsible 
for directing the animal to higher latitudes for for- 
aging, as Loggerhead Turtles are commonly encoun- 
tered in Atlantic Canadian offshore waters. However, 
the animal’s presence in nearshore waters off Nova 
Scotia remains puzzling. Loggerhead Turtles are rarely 
encountered in coastal areas of Atlantic Canada be- 
cause sea surface temperatures are normally at or 
below the lower thermal tolerance limits of this species. 

There was no formal attempt in eastern Canada to 
specifically promote the reporting of marine turtle 
sightings until 1998, when a broad public education 
program and toll-free reporting line were established 


JAMES, MARTIN, and DUTTON: GREEN TURTLE IN ATLANTIC CANADA 


The large 
pair of prefrontal scutes on the head (left) are characteristic of Chelonia mydas. However, overlapping costal and 
vertebral scutes and strongly serrated marginal scutes (right) are not characteristic of Chelonia mydas. 


for this purpose (Martin and James 2005). Continued 
collaboration with commercial fishers will be key to 
determining if the records reported here represent 
accidental occurrences of the Green Turtle in this 
region, or if small numbers of this species regularly 
forage along the Scotian Shelf in summer and early 
fall when inshore water temperatures are highest. 


Acknowledgments 

We thank K. Horne, A. Harnish and P. Young for 
reporting the turtles described here to the Nova Sco- 
tia Leatherback Turtle Working Group. Thanks are 
also extended to C. Harvey-Clark for treating the 
hybrid turtle at Dalhousie University’s Animal Care 
facilities and to J. Conway (Maritime Aquatic Species 
at Risk Office), K. Eckert (Wider Caribbean Sea Turtle 
network) and J. Gray (Bermuda Aquarium Museum 
and Zoo) for facilitating the transport to and subse- 
quent release of this turtle in Bermudian waters. We 
thank S. Mockford (Dalhousie University), R. Le Roux 
and E. La Casella (National Marine Fisheries Service) 
for assistance with genetic analysis and R. Myers 
(Dalhousie University) for helpful comments on earli- 
er drafts of the manuscript. We acknowledge support 
from World Wildlife Fund Canada, George Cedric 
Metcalf Charitable Foundation, Canadian Wildlife 
Federation, U.S. National Marine Fisheries Service, 
the Habitat Stewardship Program for Species at Risk, 
and the Natural Sciences and Engineering Research 
Council of Canada (scholarship to M.C.J.). This work 
was conducted under Department of Fisheries and 
Oceans License #2001-425. 


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Received 26 January 2004 
Accepted 6 December 2004 


A Snow-tracking Protocol Used to Delineate Local Lynx, Lynx 
canadensis, Distributions 


JOHN R. SQUIRES, KEVIN S. MCKELVEY, and LEONARD F. RUGGIERO 


U.S. Forest Service, Rocky Mountain Research Station, Forestry Science Laboratory, P.O. Box 8089, Missoula, Montana 59807 
USA. jsquires @fs.fed.us 


Squires, John R., Kevin S. McKelvey, and Leonard F. Ruggiero. 2004. A snow-tracking protocol used to delineate local lynx, Lynx 
canadensis, distributions. Canadian Field-Naturalist 118(4): 583-589. 


Determining Canada Lynx (Lynx canadensis) distribution is an important management need, especially at the southern extent 
of the species range where it is listed as threatened under the U. S. Endangered Species Act. We describe a systematic snow- 
track based sampling framework that provides reliable distribution data for Canada Lynx. We used computer simulations to 
evaluate protocol efficacy. Based on these simulations, the probability of detecting lynx tracks during a single visit (8 km 
transect) to a survey unit ranged from approximately 0.23 for surveys conducted only one day after snowfall, to 0.78 for surveys 
conducted 7 days after a snowfall. If the survey effort was increased to three visits, then detection probabilities increased 
substantially from 0.58 for one day after snowfall to about 0.95 for surveys conducted 7 days after a snowfall. We tested the 
protocol in the Garnet Range, Montana, where most lynx were radio-collared. We documented a total of 189 lynx tracks during 
two winters (2001-2003). Lynx distribution based on snow-track surveys was coincident with the area defined through radio 
telemetry. Additionally, we conducted snow-track surveys in areas of western Wyoming where lynx were believed present but 
scarce. We detected a total of six lynx tracks during three winters (1999-2002). In Wyoming , where lynx presence was inferred 


from a few tracks, we verified species identification by securing genetic samples (hairs from daybeds) along track-lines. 


Key Words: distribution, forest carnivore, Canada Lynx, Lynx canadensis, snow-track, Montana, Wyoming, surveys. 


Listing Canada Lynx (Lynx canadensis) as “Threat- 
ened” in the U. S. under the Endangered Species Act 
compels land managers to consult with the U.S. Fish 
and Wildlife Service regarding potential impacts of 
management actions (Federal Register, Volume 63, 
Number 130). One of the most fundamental informa- 
tion needs is to delineate the current distribution of 
lynx in the contiguous United States. Definitive range 
determination for rare and elusive carnivores, like the 
lynx, is difficult and may require several well-tested 
survey methods. The National Lynx Survey uses hair- 
collection pads (McDaniel et al. 2000) as a means of 
detection over large areas including much of the north- 
ern and western portions of the United States (K. S. 
McKelvey). DNA analysis of collected hair provides 
positive species identification (Mills et al. 2000), and 
is the only tested detection method during non-snow 
periods for lynx. However, unless multiple samples are 
obtained from a site, the method cannot distinguish 
local populations from single dispersing individuals, 
an important issue for land managers. 

Snow-tracking provides a survey method for detect- 
ing lynx during the winter (Halfpenny et al. 1995; 
Zielinski and Kucera 1995). Unlike hair-pad surveys, 
snow track surveys do not require a behavioral re- 
sponse, like rubbing, for detection. Lynx also have many 
attributes that make them particularly good candidates 
for snow-track surveys: lynx from southern populations 
have large spatial-use areas (approximately 150 km? for 
males and 70 km? for females; Aubry et al. 2000); have 
high daily travel rates (Ward and Krebs 1985; Mowat 


et al. 2000); and a distinctive snow-track that facilitates 
identification (Halfpenny and Biesiot 1986; Forrest 
1988). Snow-tracking also provides anecdotal infor- 
mation, such as identification of family groups, useful 
for distinguishing local populations from dispersing 
individuals. 

Snow tracking has been used extensively to survey 
lynx and other forest carnivores (Thompson et al. 1989; 
Stephenson and Karczmarczyk 1989; Beier and Cun- 
ningham 1996; Becker et al. 1999), but, like other detec- 
tion techniques, snow-track surveys have inherent 
strengths and weaknesses. Snow track surveys can be 
conducted across extensive landscapes at reasonable 
costs, and the method is sensitive to changes in carni- 
vore occupancy. However, potential problems with 
snow-track surveys include track misidentification and 
difficulties in achieving representative surveys (Half- 
penny et al. 1995; Zielinski and Kucera 1995). For ex- 
ample, Aubry and Lewis (2003) found that unscreened 
records of Fishers (Martes pennanti) that presumably 
included many “false positives” yielded a different, 
and much more extensive, distribution when compared 
to verified records. Misidentifications can also have 
serious management impacts if the detected species 
is listed as Threatened or Endangered under the U. S. 
Endangered Species Act. 

Here, we describe and evaluate a track-based survey 
method for lynx and its potential limitations. Our goal 
was to develop a reliable and representative winter sur- 
vey method to complement existing summer surveys. 
Specifically, we needed to improve on existing snow- 


583 


track survey methodologies to ensure they are repre- 
sentative in terms of their spatial extent and habitat 
coverage, and to address problems of track misidenti- 
fication by incorporating genetic sampling (i.e., scats 
and hairs frozen in tracks and daybeds). To evaluate 
reliability we tested these methods both through com- 
puter simulation and through direct comparisons with 
radio-telemetry data in an area where most lynx were 
radio-collared. 


Study Areas 

We conducted lynx surveys in the Garnet Mountain 
Range of western Montana, and in the Wyoming/Salt 
River Mountains approximately 80 km southwest of 
Jackson, Wyoming. Approximately 80% of the Garnet 
Range is forested within the Douglas-fir (Pseudotsuga 
menziesii) and Subalpine Fir (Abies lasiocarpa) series 
of the Montana Forest Habitat type (Pfister et al. 1977; 
Burcham et al. 1999). Dominant tree species include 
Subalpine Fir, Engelmann Spruce (Picea engelmannii), 
Lodgepole Pine (Pinus contorta), and Douglas-fir. Ele- 
vations range from 1160 to 2090 m. The primary use of 
forested public and private lands is timber management, 
and much of the area was extensively logged within the 
past 25 years. The Garnet Range is adjacent to private 
lands that consist mainly of irrigated hay fields and 
Big Sage Brush (Artemisia tridentata) — wheat grass 
(Agropyron spp.) cover types. 

The Wyoming and Salt River ranges, located in west- 
central Wyoming, support mixed conifer forests that 
include Subalpine Fir, Engelmann Spruce, Lodgepole 
Pine, Whitebark Pine (Pinus albicaulis), Douglas-fir, 
and Quaking Aspen (Populus tremuloides). Mesic sites 
are dominated by Subalpine fir and Engelmann Spruce 
forests in seasonally moist or wet areas, or where sub- 
irrigation maintains a high water table. Spruce-fir forests 
are typically co-dominant with Lodgepole Pine. Dry 
sites are dominated by Lodgepole Pine forests, often 
intermixed with Douglas-fir and Quaking Aspen. Pre- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


cipitation is mostly in the form of snow; elevations 
range from 1981 — 3353 m. 


Methods 
Modeling Track Detections 

We estimated the probability of detection relative 
to search effort using computer simulation to model 
track detections based on the tortuosity of actual lynx 
tracks. As part of our research, we __ back- 
tracked 15 radio-collared lynx and mapped their tracks 
using a Trimble GEO Explorer II with data points 
taken at 2 s intervals. We randomly selected one 3-km 
track from each lynx and calculated the average track 
tortuosity by dividing total length by the linear dis- 
tance traveled (Turchin 1998). This result was used 
when generating simulated tracks to ensure realistic 
track tortuosity. 

Computer simulations required that we estimate daily 
travel distances of lynx. Daily movements as meas- 
ured by radio telemetry are shorter than the animal’s 
actual movement because of unrecorded tortuosity. 
In general, reported daily movements of lynx ranged 
from 1-9 km as measured using various methods, 
with most estimates from 1-4 km (Table 1). For mod- 
eling, we assumed that daily movements varied ran- 
domly from 1.0-4.0 km (z= 2.5 km). We chose a con- 
servative estimate of daily movements to better ensure 
that our model results addressed locating lynx at low 
densities where tracks were uncommon. A uniformly- 
distributed random number was drawn from this range 
for each trial. A simulated day’s movement was com- 
pleted when the linear distance traveled between the 
day’s start and the current location exceeded the drawn 
random number. The program continued in this man- 
ner counting the number of days elapsed until the 
survey transect intersected a simulated track, and the 
number of elapsed days was written to an output file. 
Simulated tracks began at random locations within 
hypothetical, circular home ranges of 100 km? with 


TABLE |. Reported daily distances for lynx movement; only winter movements were included where both summer and win- 


ter movements were reported. 


Study Distance (km) 
Parker et al. 1983 8.8, male 
7.6, female 


6.5, juvenile 
Ward and Krebs 1985 


2.7 (95% CI = 1.8-3.7), hares common 


Method 


Cumulative distance based on 2 hour relocations 


Daily telemetry locations 


2.4 (95% CI = 2.0-2.9), hares uncommon 


Brittell et al. 1989 1.0, (range = 0.02-7.4), males 
0.7, (range = 0.02-4.0), females 
3.8 (sd = 0.6), male 


3.0 (sd = 0.4) female 


Apps 2000 


Mowat et al. 2000 
Squires and Laurion 2000 


5-9 reported average of multiple studies 
2.8 (range = 2.5-3.3), 4 Montana males 


Daily telemetry locations 
Daily telemetry locations 


Snow tracking 
Daily telemetry locations 


3.2 (range = 2.5-3.9), 3 Montana females 
2.7 (range = 0.7-9.5), Wyoming male 
2.2 (range = 0.3-5.2), Wyoming female 


2004 


an initial random direction. The length of simulated 
tracks was evaluated using travel distances that would 
be expected if we assumed that 1, 3, or 7 days had 
elapsed since the most recent snowfall. 

The survey route was modeled as a straight, 8-km 
transect, randomly located within the 100 km? home 
range. The entire transect was incorporated into the 
home range, an assumption based on the study design 
in which the simulated survey track would be repli- 
cated within each cell of a contiguous grid. Hence, at 
least 8 linear km of survey track would be found in 
any arbitrary 100 km? circular area within the survey. 


Snow-track Protocol 

We used GIS to establish an 8 x 8 km grid across 
each survey area to define sample units. The size of 
this sample unit was slightly smaller than a typical 
female home range (Aubry et al. 2000) and was em- 
ployed to reduce the chance of missing individuals, 
but was still efficient for searching large landscapes 
(Zielinski et al. 1995). Female lynx typically have small- 
er home ranges than males (Koehler 1990; Poole 1994; 
Slough and Mowat 1996). Establishing a grid of sur- 
vey units covering the study areas ensured that our 
search effort was spatially well distributed across avail- 
able habitats (Zielinski et al. 1995). After establishing 
the sample grids, we assigned a high or low priority to 
sampling units based on dominant habitat-type. Sam- 
ple units in moist, Subalpine Fir forests were a high 
priority because these stands are more heavily used by 
lynx than drier forest types (McKelvey et al. 2000). 
Our protocol specified that “high” priority 
units be surveyed at least twice per winter (Halfpenny 
et al. 1995). Open-forest types, such as dry Lodgepole 
Pine, Quaking Aspen, and Ponderosa Pine (Pinus pon- 
derosa), were “low” priority and were only searched 
once per winter. Open habitat types (large park lands, 
sage brush, tundra, agriculture) and high elevation sites 
(tundra, rock, and ice) are rarely used by lynx (Koehler 
1990; Aubry et al. 2000; McKelvey et al. 2000) and 
were not surveyed. 

Observers primarily searched for animal tracks by 
traveling 15-20 km/hr on roads and trails using snow- 
mobiles. We saw no evidence that lynx were reluctant 
to cross snowed-in logging roads and trails (Squires; 
unpublished data; see also O’Donoghue et al. 1998). 
Survey units that could not be surveyed by snowmobile 
were searched on snowshoes to the extent possible. 
The protocol specified that we search 10 km in each 
8 x 8 km sample unit choosing routes that preferen- 
tially traversed forested habitats with high horizontal 
cover. This search effort per sample unit was sufficient 
to traverse key areas within the unit, but still allowed 
us to efficiently search multiple sample units per day. 
High horizontal cover is an important component of 
Snowshoe Hare (Lepus americanus) habitat, the domi- 
nant prey species of lynx (Hodges 2000). Sample units 
that contained only dry, open forests were surveyed 
using routes that best bisected the search area. 


SQUIRES, MCKELVEY, and RUGGIERO: SNOW-TRACKING PROTOCOL 


585 


Our goal was a representative survey of sample units 
that were spatially well distributed across available 
habitats, while preferentially searching the best habitats 
in each unit to maximize the chance of detecting lynx. 

We recorded the locations of all lynx tracks and the 
tracks of other carnivores using a Trimble GEO Ex- 
plorer Il GPS. Multiple crossings known to be the same 
animal and <100 m apart were recorded as a single 
detection. We measured (stride, straddle, length, width, 
depth; Halfpenny et al. 1995) and photographed all rare 
carnivore tracks to provide documentation. Track mis- 
identification is a potential problem of snow-based 
surveys (Zielinski and Kucera 1995; Halfpenny et al. 
1995), especially in areas where the species of interest 
is rare. We collected genetic samples (e.g., hairs, scats) 
from snow-tracks on the Wyoming study area to pro- 
vide unequivocal documentation. To collect genetic 
samples, we followed presumed lynx tracks in search 
of scats or day beds. We carefully removed loose snow 
from daybeds until we reached the frozen bed layer. 
We then thoroughly searched the snow surface and used 
tweezers to secure hairs frozen in the snow. Hairs were 
stored in desiccant until the Rocky Mountain Research 
Station’s Wildlife Genetics Laboratory, Missoula, Mon- 
tana, extracted and analyzed DNA samples. 

Snow-track surveys require trained personnel to be 
effective (Zielinski and Kucera 1995). We trained ob- 
servers at our long-term lynx study area near Seeley 
Lake, Montana. Observers back-tracked radio-collared 
lynx to practice identifying tracks under diverse envi- 
ronmental conditions (e.g., forest type, snow conditions, 
topography). Following this training, we assumed that 
all observers could recognize lynx tracks. 


Survey Verification 

In the Garnet Range, we used radio-telemetry data to 
determine if the distribution of lynx within the study 
area was coincident with survey results. Lynx were 
trapped using specially designed box traps baited with 
carrion, beaver castor, and pie plates and wings for 
visual lures (Kolbe et al. 2003). Trapping was conduct- 
ed throughout the winters of 2001-02 and 2002-03. 
Lynx were chemically immobilized using a syringe 
pole to administer a mixture of ketamine (10 mg/kg 
Ketaset®; concentration 100 mg/mL) and xylazine 
(1 mg/kg; concentration 100 mg/mL); this dose pro- 
duced predictable immobilization periods (30-40 min- 
utes) and stable vital signs. We fitted adult lynx with 
VHF radio transmitters (170 g), weighed, measured, 
and extracted blood from each for future DNA analy- 
sis. Kittens were fitted with a padded, 100 g collar that 
allowed for growth. We monitored lynx movements 
using aerial telemetry (90% of locations) approximately 
twice per month augmented with ground-based tel- 
emetry. 

We combined all relocation points and calculated a 
95% kernel home range to delineate the distribution of 
the local lynx population based on telemetry (Worton 
1989; Seaman et al. 1999). We then overlaid a 95% 


586 


fixed kernel home range on the distribution of lynx, 
as delineated by snow tracking, to determine if the 
two methods yielded coincident distributions. 


Results 
Simulations of track detections 

Computer simulations of track detections indicated 
different asymptotic relationships between the proba- 
bility of detection and survey effort (number of visits 
per winter), based on the time between snowfalls suf- 
ficient to obliterate tracks (Figure 1). The probability 
of detecting lynx tracks during a single visit (8 km 
transect) to a survey unit ranged from approximately 
0.23 for surveys conducted only one day after snow- 
fall, to 0.78 for surveys conducted 7 days after a snow- 
fall. If the survey effort was increased to three visits, 
then detection probabilities increased substantially from 
0.58 for one day after snowfall to about 0.95 for surveys 
conducted 7 days after a snowfall. Even a modest sur- 
vey effort of two visits would detect tracks approxi- 
mately 80% of the time, provided more than 3 days 
had passed since the last snowfall. 


Field test — Garnet Range, Montana 

During winters 2001-2002 and 2002-2003, we sur- 
veyed for lynx on 242 km and 438 km of roads and 
trails, respectively. We documented 37 track detections 
of lynx in 4 of 12 survey units during winter 2001- 
2002, and 37 detections in 4 of 16 survey units during 
winter 2002-2003. In addition, we observed 115 lynx 
tracks during winter 2001-2002 and 53 in 2002-2003 
in survey units, but not during the formal survey. These 
detections were made during trapping and related trav- 
el. We encountered a higher number of track-cross- 
ings during trapping because we only trapped in 
areas of high lynx use. The average elevation where we 
located lynx tracks was 1839 m (SD 97, range 1443 — 
2012 m). 

We trapped five lynx during winter 2001-2002, and 
two additional individuals during winter 2002-2003. 
We believed this sample of radio-collared individuals 
was a local census of lynx in the Garnet Range based 
on trapping results, recaptures, and field observations, 
but we could not formally test this perception. 


Field test — Wyoming and Salt River Ranges, Wyoming 

We conducted track-surveys for lynx in Wyoming 
during the winters of 1999-2000, 2000-2001, 2001- 
2002. During winter 1999-2000, we surveyed 1055 km 
in 32 sample units, and we detected one lynx track in 
the Salt River Range and a second track in the north- 
eastern portion of the Wyoming Range. In addition, 
tracks from a family group were observed in the north- 
eastern portion of the Wyoming Range by Wyoming 
Game and Fish Department biologists (B. Oakleaf, 
personal communication 2000). During winter 2000- 
2001, we surveyed 1103 km in 37 sample units, and 
4 lynx tracks were detected in the northeastern portion 
of the Wyoming Range, and one set in the southwestern 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


© 
fo) 


Probability of detecting lynx 
oO 
(op) 


Number of visits 


FIGURE |. Computer-modeled relationship between the proba- 
bility of detecting lynx and the number of visits to an 8 km 
survey transect pixel relative to the number of days since 
last snow. 


Salt River Range. We surveyed 1080 km in 34 sample 
units during winter 2001-2002 and detected no lynx. 

Two tracks located during the 1999-2000 survey 
looked like lynx, but were poorly defined with each 
impression being a pedestal above the snow surface 
as a result of wind scouring. We followed two tracks 
for approximately 200 m until we located daybeds and 
collected five hairs. Two of the hairs contained suffi- 
cient DNA to amplify and the species identification 
was verified as lynx. 


Survey Verification 

The distribution of lynx in the Garnet Range, as de- 
fined by a 95% fixed kernel home range of all marked 
animals (n= 96 locations from 7 individuals, = 14 loca- 
tions per individual, Figure 2B) combined, subsumed 
97% of all tracks detected during snow-track surveys. 
The general distribution of lynx in the Garnet Range 
as defined by radio telemetry and snow-track surveys 
was coincident (Figure 2A). 

We documented too few lynx detections in the Wy- 
oming and Salt River ranges to delineate a distribution 
based on snow-track surveys. However, biologists with 
the Wyoming Game and Fish Department trapped two 
lynx in the northeastern portion of the Wyoming Range 
during the winter of 1996-1997 (B. Oakleaf personal 
communication 2000). The winter home ranges of these 
two individuals were restricted to the northeastern por- 
tion of the Wyoming Range in the same approximate 
area as 5 of the 7 track detections from our survey. 


Discussion 

Snow-tracking has been used to document the pres- 
ence/absence of carnivores (Zielinski and Kucera 1995), 
and continues to be an important technique, especially 
when the organisms of interest are rare and difficult 
to observe. One primary appeal of using track-based 
surveys for presence/absence sampling is the method’s 


2004 SQUIRES, MCKELVEY, and RUGGIERO: SNOW-TRACKING PROTOCOL 587 


- # di 
a 
. 
Ke f j 
eS er! rae 
SS re 
\ i) a 
as / ; 
| oe 
| me, 
| XX 
—, 
) - a 
1 &Pevil Mn 
} 4% : q ; d ¥ fs 
f y © Lynx Telemetry Locations s Drummond 
es 95% Fixed Kernal Home Range << 
acs velude Grid 


4 a Lynx Detections 
RASS Snow-track Surveys 


f ia = Lynx Distribution —_ Drummond 


Survey Grid 


i | 
| 95% Fixed Kernal Home Range (Telemetry - See Above) 


Sa a 
FicurE 2. The local distribution of lynx in the Garnet Range delineated from: (A) radio telemetry of all individuals com- 


bined (95% fixed kernal, n = 96 locations from 7 individuals, x = 13 locations per individual) compared to, (B) the local 
distribution based on sample units with track detections. 


588 


sensitivity to changes in use patterns, especially for 
species like lynx that are highly mobile and have large 
home ranges (Ward and Krebs 1985; Poole 1994; Slough 
and Mowat 1996; Aubry et al. 2000). Changes in lynx 
occupancy can be quickly detected across extensive 
areas using snow-track surveys. For example, general 
occurrence data from 1842-1998 suggested that lynx 
had a long-term history of occupancy in the Salt River 
and Wyoming Ranges (see McKelvey et al. 2000: Fig- 
ure 8.17), and the perception that lynx were currently 
distributed throughout the Wyoming and Salt River 
Ranges was shared by local managers. Snow-track sur- 
veys conducted in a representative manner throughout 
the Wyoming and Salt River Mountain ranges provided 
evidence that lynx were present in 1999, and this result 
was confirmed using genetic analyses in 2000. How- 
ever, the low overall detection rate coupled with the lack 
of tracks in 2001 indicated that lynx were rare and 
only located in a small area in the northern part of the 
Wyoming range. 

In areas with many track detections, snow-track sur- 
veys can yield more than simple presence/absence data. 
Results from the Garnet Range demonstrated that snow- 
track surveys conducted in a representative manner can 
also be used to delineate the local distribution of lynx. 
The high overlap in distributions determined through 
telemetry and survey data suggested that snow-track 
surveys can at least coarsely define the local distribu- 
tion of lynx (Figure 2). We acknowledge that survey data 
are more limited than telemetry data given that only 
winter movements are documented and the resulting 
distribution is coarse-scale relative to the sample unit. 
However, even a coarse-scale understanding of local 
distributions is useful to habitat managers. Thus, based 
on our data in the Garnet Range, we believe that snow- 
track surveys that are spatially well distributed across 
all available habitats can be used to estimate the local 
distribution of lynx during the winter. 

If lynx are abundant in an area, a 5% misidentification 
rate would have little impact on management decisions. 
However, in areas where lynx are very rare, a track 
misidentification can have serious management rami- 
fications. One way to address the chronic problem of 
track misidentification is to view the track as a “‘col- 
lection device” for obtaining genetic samples rather 
than a primary method of species identification. Genet- 
ic verification provides definitive species documentation, 
regardless of the observer’s ability to identify rare car- 
nivores based on track characteristics. We therefore rec- 
ommend that genetic identification be included when- 
ever proper identification of an individual set of tracks 
is critical. 

Snow-track surveys are labor intensive, and we have 
only used this method at a mid or “meso” spatial scale, 
where we searched all sample units. Such an intensive 
approach would be impractical across large landscapes, 
for example a large portion of a state. Although we 
have not conducted a snow-track based survey at a 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


“state” scale, adaptive cluster sampling may provide 
a promising method of achieving a representative sam- 
ple across very large landscapes (Thompson 1992). 
Adaptive cluster sampling is effective for delineating 
clumped distributions, which we believe is the case for 
lynx at the southern extent of the species’ range (Aubry 
et al. 2000). 

Snow-track surveys are often conducted when track 
characteristics are most distinctive, usually within a 
day or two following fresh snowfall. This constraint 
limits the utility of snow tracking because the proba- 
bility of detecting lynx and other rare carnivores is a 
function of the time since last snow. The more time 
that lynx have to travel and establish long tracks, the 
more likely it is that their tracks will be detected on a 
survey route. For example, our simulations suggest 
almost a 100% probability of detecting lynx in two 
entries 7 days after a snowfall compared to a 43% 
probability of detection if the area was searched 1 day 
from snowfall (Figure 1). In the areas we surveyed, snow 
accumulations over 5 cm were recorded on 22 and 24 
(17-18%) of the days for the Garnet and Wyoming 
ranges, respectively, and the snow-free periods 
between storms varied from 4-6 days (U.S.D.A., Nat- 
ural Resources Conservation Service, SNOTEL data). 
The ability to correctly identify “old” tracks by incor- 
porating genetic samples allows biologists to survey 
for rare carnivores when the probability of detection 
is high, even under poor tracking conditions. 

The winter surveys that we used to detect lynx in 
Wyoming and Montana had the following key ele- 
ments: (1) sample units were representative in terms 
of spatial coverage, but still focused on high-quality 
habitats that maximize lynx detections; (2) search 
effort per survey grid was defined and consistent across 
the survey area; (3) search intensity was consistent 
with simulated results indicating a high probability 
of detection, (4) technicians were well trained in track 
identification and data recording; (5) track identifica- 
tions were rigorously documented and spatially ref- 
erenced; and (6) lynx tracks were searched for genetic 
samples to confirm identification. We believe that sur- 
veys having these characteristics can be used to rigor- 
ously define the range of local lynx populations, and 
could be extended to the regional scale using adap- 
tive cluster sampling. 


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Received 4 February 2004 
Accepted 17 August 2004 


Hummock Vegetation at the Arctic Tree-line near Churchill, Manitoba 


JORG TEWS 


Institute of Geography, University of Erlangen-Niirnberg, Kochstr. 4/4, 91054 Erlangen, Germany 
Current address: Geomatics and Landscape Ecology Research Laboratory (GLEL), Department of Biology, Carleton University, 
1125 Colonel By Drive, Ottawa, Ontario K1S 5B6 Canada 


Tews, Jérg. 2004. Hummock vegetation at the arctic tree-line near Churchill, Manitoba, Canadian Field-Naturalist 118(4): 
590-594. 


Hummocks, small earth or peat mounds, are widely distributed in the arctic and develop as a consequence of biomass accumulation 
and cryoturbation in the active layer. There is general agreement that the type of vegetation covering peat hummocks may 
alter the accumulation rate of organic material and thus hummock growth and local carbon sink dynamics. Studies on hummock 
plant community compositions from the arctic are very scarce. Here, I present results of a case study from the arctic tree-line 
near Churchill, Manitoba (Canada). Vegetation composition, hummock height and soil moisture content were recorded in 
40 peat hummocks located along a tree-line gradient from open forest to tundra. Based on a cluster analysis I found three 
moss-dominated types of hummock vegetation, according to (1) a Tomenthypnum nitens (golden fuzzy fen moss) type on 
low hummocks, (2) a Hylocomium splendens (stair-step moss) type on medium-sized hummocks, and (3) a Pleurozium schreberi 
(red-stemmed feathermoss) type on hummocks higher than 60-70 cm. I found hummock height to increase towards the forest 
interior with decreasing water content of the upper organic layer on the hummock top. This is indicated by a significant change 
in vegetation composition towards drought resistant moss species on higher hummocks. Furthermore, species richness decreased 
with increase in hummock height. Based on evidence from historical tree-line invasion the overall results suggest that hammock 
height increases due to peat accumulation over the course of time resulting in a typical change in plant community composition. 


Key Words: peat hummocks, Tomenthypnum nitens, Golden Fuzzy Fen Moss, Hylocomium splendens, Stair-step Moss, Pleu- 


rozium schreberi, Red-stemmed Feathermoss, subarctic, forest-tundra ecotone, Hudson Bay Lowlands, Manitoba. 


Hummocks are small, up to 1 m high soil mounds 
widely distributed in the northern boreal, sub-arctic 
and arctic permafrost regions (e.g., Lundquist 1969; 
Mackay 1980). Generally, two types of hummocks can 
be classified: earth hummocks and peat hummocks. 
Earth hummocks may develop as a result of frost heave 
and cryoturbation processes where the organic layer 
overlies fine-grained frost-susceptible soils (Quinton 
et al. 2000). In contrast, peat hummocks grow as the 
result of the accumulation of organic material and 
where the surface of the uppermost mineral layer inside 
the hummock is positioned below the surrounding 
ground level (see Dredge 1992). Both hummock types 
may be perennially frozen or, as a result of warmer 
summer climate and lower latitude, completely thawed 
(Zoltai and Pettapiece 1974; Tarnocai and Zoltai 1978). 
As a transitional form they may be partially frozen with 
ice lenses remaining in the hummock core during the 
summer period. 

Even though hummocks are a common landscape 
feature and hummocky terrain covers a large proportion 
of Arctic Canada (Tarnocai and Zoltai 1978), scientific 
studies focusing on the vegetation structure of peat 
hummocks are very scarce. This lack of empirical data 
and also the potential role of peat hummocks for the 
carbon dynamics in the northern hemisphere empha- 
size the need to investigate the spatial distribution of 
major hummock plant community types. Besides the 
prevailing local climate, carbon storage in the arctic 


is strongly influenced by vegetation composition and 
succession (see Camill et al. 2001). Thus, in order to 
appreciate carbon dynamics in the northern hemisphere 
acknowledge of potential carbon sinks (such as peat 
hummocks) and their major plant community types are 
necessary. This enables evaluation of ecological factors 
which may alter accumulation or decomposition rates 
of organic material and related carbon dynamics. For 
example, mosses which often dominate hummock veg- 
etation have the potential to play a key role in modifying 
decomposition rates and the thermal and hydrological 
regime of arctic soils (Beringer et al. 2001). Here, I 
present results of a case study on vegetation composi- 
tion, succession and physiognomy of 40 peat hum- 
mocks located along an arctic tree-line transect from 
tundra to open forest in the Hudson Bay Lowlands 
near Churchill, Manitoba (Canada). 


Methods 
Study area 

The study site is situated at the open forest tree-line 
near Twin Lakes, a flat-topped glacial kame deposit 
approximately 25 km southeast of the town of 
Churchill (Figure 1). The open forest vegetation is com- 
posed of a mix of Tamarack (Larix laricina [Du Roi] 
K. Kock) and White Spruce (Picea glauca [Moench] 
Voss) with interspersed Black Spruce (Picea mariana 
[Mill.] Britt Sterns & Pogg). The present tree line 
north of Twin Lakes is extended into a wet sedge fen 


590 


2004 


roy 


udson|Bayss 


S 
= 
es 
SiS 
apa 
ov 


Twin Lakes 


FiGuRE |: Location of the study site in the Hudson Bay 
Lowlands near Churchill in northeastern Manitoba 
(Canada). 


predominated by Carex aquatilis (Water Sedge) and 
Carex limosa (Mud Sedge). The current position of the 
tree-line has moved up to 150 m towards the open 


_ sedge fen within the last 70 years (see Scott et al. 1987). 
_ Here, the current tree-line is composed of L. laricina 


which established during the latest forest invasion. 


_ Whereas young L. laricina tree-line stands are char- 


acterized by small hummocks, the open forest interior 
towards Twin Lakes is dominated by mature P. glauca 
trees on typically hummocky terrain with large hum- 


_ mocks and inter-hummock, water-filled troughs. The 


nomenclature for plants follows after Porsild and 
Cody (1980). 


_ Sample design 


During July 1999 I established a 250 m transect from 
the open sedge fen towards the forest interior. I sam- 
pled each hummock (total 40) that occurred within a 
5 m wide corridor along the transect line. For each 
hummock I measured its height from the top to the 
base. Vegetation composition was studied by using a 


_ frame placed on top of each hummock. The frame size 
0.5 * 0.5 m was small enough to cover the tops of the 


smallest hummocks. For larger hummocks I chose 
homogeneous parts of the vegetation on the hammock 
top. Vascular plant cover within the sampling frame 
was then estimated for each plant species separately 


_ using the decimal Londo-scale (Londo 1984). Within 


each frame soil samples were taken from the upper 
organic layer (5 cm — 20 cm depth) during one sam- 
pling day and then water content gravimetrically deter- 
mined (samples were 24h oven-dried at 95°). 


Statistical analysis 

To group species datasets Mulva’s minimal variance 
clustering technique using van der Maarel’s coefficient 
was used (Wildi and Orléci 1996). Simple linear regres- 
sion was used to examine the relationship between 
hummock height as an independent variable and water 
content, species richness, and similarity of species com- 


TEWS: HUMMOCK VEGETATION AT THE ARCTIC TREE-LINE 


59] 


position (species turnover rate) as the dependent vari- 
ables. For the species turnover rate between sample i 
and j hummocks were ranked by height and / defined 
according to: 


B= 


where / is the number of species that disappeared 
between sample i and j and g the number of new species. 


l+g 
i+] 


eqn |, 


Results 

Hummocks are a dominant micro-topographical fea- 
ture at the open forest tree-line. For the transect area I 
found a mean density of 320 hummocks ha’!. However, 
density was significantly higher in mature P. glauca 
stands towards the open forest. The majority of hum- 
mocks were predominately covered by either one of 
the moss species Tomenthypnum nitens (Golden Fuzzy 
Fen Moss), Hylocomium splendens (Stair-step Moss) 
or Pleurozium schreber (Red-stemmed Feathermoss) 
whereas herbaceous plants where less frequent. This 
was confirmed by a resemblance matrix of a cluster 
analysis for 40 sample plots (Figure 2). The Tomen- 
thypnum nitens-group was mainly composed of Poly- 
gonum viviparum (Alpine Bistort), Eguisetum varie- 
gatum (Variegated Scouring-rush), Andromeda polifolia 
(Dwarf Bog-rosemary), and Carex aquatilis (Water 
Sedge), indicating somewhat wet conditions (see Table 
1). In contrast, hammocks with Hylocomium splendens 
had a more or less Dwarf Shrub dominated cover with 
species such as Vaccinium vitis-idaea (Lingonberry), 
Ledum groenlandicum (Common Labrador Tea), and 
the lichen Cladina rangiferina (Grey Reindeer Lichen). 
The third major group was dominated by Pleurozium 
schreberi associated with Betula glandulosa (Dwarf 
Birch) and the grass Calamagrostis canadensis (Blue 
Joint), indicating somewhat dry conditions. 

The species groups that were revealed by the clus- 
ter analysis showed significant affiliation in terms of 
transect position and respective hummock height. The 
Tomenthypnum nitens-group was mainly found on low 
hummocks (see Table 1) located near the tree-line (Fig- 
ure 3a). Here, the organic layer on hummock tops was 
mostly saturated (Figure 3b), typical for the hydro- 
logical situation near the sedge fen. Hummocks with 
Hylocomium splendens had medium-sized heights 
and an intermediate transect position, whereas Pleu- 
rozium schreberi-hummocks with heights above 60 cm 
where predominately found in the open forest interior. 

Overall, there was an increase in hummock height 
towards the open forest (Figure 3a, R? = 0.57). Increase 
in height had a significant negative effect on organic 
layer water content (Figure 3b, R*= 0.46). In terms of 
patterns of plant species richness, species number de- 
creased with increase in hummock height (Figure 3c, 
R? = 0.42; see also Table 1). Total number of plant 
species found, including mosses and lichens, was 45, 
and 8.1 species per sample plot on average. More- 
over, intermediate hummock heights were indicated 


592 


-4.66E-02 


4.66E-01 


9.79E-01 
31 


26 

22 

34 

28 oa 

27 
Tomenthypnum 20 


nitens - 
Gruppe 21 


Pleurozium 
schreberi - 
Gruppe 


Hylocomium 
splendens - 
Gruppe 


a ee | ee eee] 
9.79E-01 4.66E-01 -4.66E-02 


FiGuRE 2: Cluster analysis of 40 hummock vegetation sam- 
ples. Three moss-dominated plant community types are 
distinctive according to the occurrence of Tomenthyp- 
num nitens, Hylocomium splendens and Pleurozium 
schreberi. Numbers for each sample indicate respec- 
tive hummock height given in cm. 


by a relatively high species turnover rate when plots 
were ranked by height (Figure 3d), i.e., species com- 
position showed a higher variation than vegetation of 
either low or high hummocks. 


Discussion 

The results of this field study from the arctic tree- 
line near Churchill indicate a significant relationship 
between hummock height and the position along the 
tree-line gradient on one hand, and hummock height 
and water content, vegetation type, species richness, 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


and species turnover on the other hand. Increasing 
hummock height towards the forest interior seemed to 
reduce moisture availability for mosses on the hum- 
mock tops and facilitate the establishment of species- 
poorer communities with drought-resistant vascular 
plants. Interestingly, Zomenthypnum nitens, Hylocomi- 
um splendens and Pleurozium schreberi are ubiqui- 
tous moss species with a wide ecological distribution 
(Nicholson and Gignac 1995). However in this study 
they showed distinct distribution patterns in relation 
to hummock height and soil water availability within 
a relatively small area. 

The plant community composition of peat hammocks 
described here are the first inventory of hummock 
vegetation in the Hudson Bay Lowlands, the largest 
contiguous wetland in North America (Boudreau and 
Rouse 1995). Other published studies are concerned 
with the Mackenzie delta region where earth hummock 
physiology and plant species composition is com- 
pletely different. Thus, they are difficult to compare 
(see e.g., Zoltai and Pettapiece 1974). It is unclear 
whether peat hummocks accumulate organic material 
and increase in size regardless of the local environ- 
ment or whether this is driven by the micro-topogra- 
phy such as the establishment of trees. However, the 
local tree-line extension near Twin Lakes suggests that 
once young trees establish on formerly open tundra, peat 
hummocks may develop where shading and increased 
moisture from trapped snow coincide with feather moss 
establishment (Scott and Hansell 2002). Moreover, 
the occurrence of rotten tree stumps in the subsoil of 
the former centre of large, degraded hummocks (J. Tews 
personal observation) may support the latter hypothesis 
and is additional evidence that these hummock are 
formed by organic matter accumulation, not by cryo- 
genic processes as is the case with earth hummocks. 

Based on this study I hypothesize that hummock 
plant community composition at the tree-line near Twin 
Lakes is changing in the course of hummock growth. 
This is important as accumulation rates of peat (and 
thus carbon dynamics) are strongly influenced by local 
vegetation succession (Camill et al. 2001). Bello and 
D’Souza (2000) found that with increase in hum- 
mock height, accumulation rates of organic material 
decrease. For an average height of 20 cm they estimated 
an accumulation rate of 20 g m? year’, for 60 cm hum- 
mock height only 5 g m? year". These results largely 
confirm growth rates found for Tomenthypnum nitens 
and Hylocomium splendens (see Busby et al. 1978). 
In general, decomposition rates increase with decreas- 
ing water saturation during the short summer period. 
However, due to this rather small array of data, current 
available data may not support proper estimates of 
community-specific accumulation rates necessarty to 
model large-scale spatial and temporal carbon dynam- 
ics in the arctic. Thus, based on the knowledge of the 
predominant types of hummock plant communities, 
more studies on community-specific accumulation 
rates are needed. 


| 
H 2004 TEWS: HUMMOCK VEGETATION AT THE ARCTIC TREE-LINE 593 


| Hummock height vs. gradient position Water content 
a) @ 120 b)  100,-——_____ == 
Ss ~ R“=0.462, p<0O. 
5 80 a areal 
oO 3) 
Oo 80 
2 60 ie SH Lges gs 
4 40 E ro oe 
i) 
& 20 2 60 © e 
= 1 {0) 
x 50 
0 ee Ale 0 20 40 60 80 100 120 
Gradient [m] *f a 
Tuda ————— > Open forest ummock eight [cm] 
Species richness 8 — turnover rate 
oii 
@) 4s 
a R2= 0.4209, p < 0.€ rian R* = 0.2563, p < 0.C 
8 oe FS 
8 10 206 
oa = 
= B04 
ra | 
che ca 0.2 
5 
Z, 0 0 
0 20 40 60 80 100 120 0 20 40 60 80 100 120 
Hummock height [cm] Hummock height [cm] 


| Ficure 3: Simple regression analysis for: (3a) hummock height vs. gradient position along the tree-line from open tundra to 
forest; (3b) water content of upper humus layer on hummock top vs. hummock height; (3c) total number of plant 
species vs. hammock height; (3d) species turnover rate vs. hummock height. 


Another current environmental issue makes the mat- agreement that global climate change will alter decom- 
| ter even more complex: global climate change. On __ position rates and carbon storage (e.g., Gorham 1997; 
| average, climate change in the arctic may yield more _ Earle et al. 2003). Dormann and Woodin (2002) point- 
| precipitation and warmer summers (see Sonesson ed out that the driver of future change in arctic vege- 
| 2002). Based on these assumptions, there is general _ tation is likely to be increased nutrient availability, aris- 


Table 1: Typical species composition of hummocks at the arctic tree-line near Churchill, Manitoba. The table shows 27 
characteristic hammock samples out of a total of 40 samples (13 samples with less significant community affiliation are not 


| shown). Three moss-dominated plant community types are evident according to Tomenthypnum nitens, Hylocomium splendens 
) and Pleurozium schreberi. 


Dp ies BER Sy 6s ONO AP Aa AS 14y SANG 17; 16H 19 4 (20): 21) 22) 23) 242526 27 
‘Hummock height 21 19 26 28 34 31 25 20 25 37 22 22 27 98 47 98 73 77 61 10595 76 11277 84 67 80 
# of species OMe Onno SSN Oron iO CON niOnn GR Gus GuSie ae fai nlem OMS) a9 hater Ouro 
_Tomenthypnum nitens Sei GUORMOeTOLOnaSrwtOe GiB bid OO Li29 
Polygonum viviparum Moana Ae Onn Aw Aan Ql a2 

| Equisetum variegatum Ana 2s AL 4 OE TAuL eit 4 °.2 4 

| _Scirpus caespitosus ne 3 4 4 
Carex aquatilis 1 1 
_ Andromeda polifolia 2 1 


Platanthera obtusata 
_Oxyecocus microcarpus 
Rubus acaulis 
_ Pedicularis lapponica 
Salix reticulata _ 
Aulacomnium palustre 


Hylocomium splendens i 


iS 
_Vaccinium vitis-idaea CEC, Bey Abi 40 Wig ln eg! 
Ledum groenlandicum 4.4 2: 
_Ledum decumbens 2 ES TY 1 
_Cladina rangiferina. c4iuat i PY year sar 
Pleurozium schreberi (4 StS OS 


_ Betula glandulosa 
Calamagrostis canadensis 


io 
aA 


594 


ing for example from temperature-induced increases 
in mineralization. In particular, the response of plant 
growth to rising CO, levels appears to depend on nutri- 
ent availability (Heijmans et al. 2002). In the arctic 
vegetation nitrogen is tightly controlled by the moss 
layer (e.g., Li and Vitt 1997; Sommerkorn et al. 1999). 
Moreover, mosses such as Pleurozium schreberi have 
been reported to be able to fix nitrogen in symbiosis 
with a cyanobacterium (i.e. Nostoc sp.) (Deluca et al. 
2002). Thus, increase in average summer temperatures 
due to climate change and the resulting increase in min- 
eralization rates may potentially decrease local carbon 
storage with severe consequences for the global carbon 
cycle. However, scientific research is far from estab- 
lishing realistic future scenarios. For example, in the 
context of our study, it was recently shown that indi- 
vidual bryophyte species displayed contrasting respons- 
es to changes in the nutrient supply and that they should 
not be grouped as a single functional type (Gordon et 
al. 2001). From this it is evident that further detailed 
hummock vegetation studies are needed, particularly 
in the arctic where the impact of climate change is 
likely to be most effective. 


Acknowledgments 

I thank the Churchill Northern Studies Centre and 
the Northern Research Fund (NRF) for logistical and 
financial support. Special thanks go to C. Oswald and 
P. Brown for their great help during the field work. I 
appreciate the constructive comments on this manu- 
script by two anonymous reviewers. 


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Received 17 February 2004 
Accepted 24 December 2004 


Effects of Lodgepole Pine Dwarf Mistletoe, Arceuthobium americanum, 
on Jack Pine, Pinus banksiana, Growth in Manitoba 


_ Brock Epp and JACQUES C. TARDIF'! 


| Centre for Forest Interdisciplinary Research (C-FIR) and Department of Biology, University of Winnipeg, 515 Avenue 
Portage, Winnipeg, Manitoba R3B 2E9 Canada 
‘Corresponding author 


| Epp, Brock, and Jacques C. Tardif. 2004. Effects of Lodgepole Pine Dwarf Mistletoe, Arceuthobium americanum, on Jack 
Pine, Pinus banksiana, growth in Manitoba. Canadian Field-Naturalist 118(4): 595-601. 


| The Lodgepole Pine Dwarf Mistletoe (Arceuthobium americanum Nutt. ex Engelm.) is an important pathogen of Jack Pine 
| (Pinus banksiana Lamb.). Dwarf Mistletoe alters tree form, suppresses growth, and reduces volume and overall wood quality 
of its host. Stem analysis and a 3-parameter logistic regression model were used to compare the growth of heavily and lightly 
to non infected Jack Pine trees. At the time of sampling, no significant reduction in diameter at breast height and basal area 
were observed in heavily infected trees. However, a significant reduction in height and volume and an increase in taper were 
observed in heavily infected trees. Growth models predicted a 21.1% lower basal area, 23.4% lower height and 42.1% lower 
volume by age 60 for the high infection group. 


Key Words: Lodgepole Pine Dwarf Mistletoe, Arceuthobium americanum, Jack Pine, Pinus banksiana, stem analysis, logistic 
regression, basal area, height, volume, tree growth, productivity, Manitoba. 


Le Faux—gui du pin (Arceuthobium americanum Nutt. ex Engelm.) est un important agent pathogeéne du Pin gris (Pinus banksiana 
Lamb.). Le faux—gui modifie le défilement des arbres hotes, réduit leur croissance et leur volume marchand ainsi que la qualité 
du bois. L’analyse de tige et la régression logistique a trois-paramétres ont été utilisés afin de comparer |’accroissement 
entre les Pins Gris sévérement atteints et faiblement atteints. Au moment de |’ échantillonnage, aucune réduction significative 
du diamétre a hauteur de poitrine ou de la surface terriére ne fut observée chez le groupe sévérement atteint. Toutefois, les 
arbres atteints ont enregistré une baisse significative de la hauteur et du volume ainsi que du coefficient de défilement. Les 
modeéles de régression ont prédit une perte de 21,1% en surface terriére, de 23,4% en hauteur et de 42,1% en volume pour 


les pins gris fortement atteints par le faux—gui a un age de 60 ans. 


Mots clés: Le Faux—gui du pin, Arceuthobium americanum, Pin gris, Pinus banksiana Lamb., analyse de tige, régression 
logistique, surface terriére, hauteur, volume, croisssance, productivité, Manitoba. 


Jack Pine (Pinus banksiana Lamb.) is an important 
commercial tree species because of its pole-like growth 
form, and as a pioneer species, it is relatively easy to 
regenerate following harvest (Rudolph and Laidly 1990; 
Sims et al. 1990). An important pathogen of Jack Pine 
is Lodgepole Pine Dwarf Mistletoe (Arceuthobium 
americanum Nutt. ex Engelm.). There are five species 
of Dwarf Mistletoe known in Canada, of which A. 
americanum is the most widely distributed, extend- 
ing from British Columbia to southeastern Manitoba 
(Hawksworth and Wiens 1996). Individual species of 
Dwarf Mistletoe are generally host-specific, able to 
infect only a few tree species. In the case of A. amer- 
icanum, other susceptible hosts include Lodgepole 
Pine (Pinus contorta Doug}. ex Loud.) and Ponderosa 
Pine (Pinus ponderosa Dougl. ex P. & C. Laws.). 

As of 1996, there were 670 000 ha of forest with 
severe A. americanum infections in Alberta, Saskat- 
chewan and Manitoba (Brandt et al. 1998). In Mani- 
toba, approximately 8.7% of mature Jack Pine stands 
in important growing regions were infected by Dwarf 
Mistletoe (Baker et al. 1992). This has resulted in a loss 
of up to 7.9% of the total pine volume in these regions 


(525 224 m? out of 6 648 405 m°) and a loss of up to 
70.3% within the infected stands. Aside from the sub- 
stantial volume losses, there is also a reduction in wood 
quality, making wood less merchantable. Wood of 
Lodgepole Pine infected by A. americanum exhibited 
greater longitudinal shrinkage and was weaker in 
strength when compared to uninfected trees because of 
the production of tree-rings having a lower percentage 
of latewood, shorter tracheid length, and higher resin 
content (Piirto et al. 1974). Generally, Dwarf Mistletoe 
deprives its host of water and nutrients, thus reducing 
height and diameter growth as well as seed production, 
and weakening the tree (Franc and Baker 2000). Dwarf 
Mistletoe alters tree form by disrupting apical domi- 
nance (Tinnin and Knutson 1980) through formation 
of witches’ brooms. Because Dwarf Mistletoe derives 
all of its nutrients from host tissue and fixes little or 
no carbon dioxide for its own use, witches’ brooms act 
as sinks for metabolites produced in other parts of the 
host (Hull and Leonard 1964a, 1964b). Douglas-fir and 
Western Larch infected by A. douglasii and A. laricis, 
respectively, exhibited a significant increase in leaf to 
sapwood ratios in heavily infected trees, altering resource 


395 


596 


allocation processes and reducing the overall water- 
use efficiencies (Sala et al. 2001). 

The long incubation period characterized by the 
life cycle of A. americanum (Hawksworth and Wiens 
1996) poses a problem in the early detection of initial 
infections. In forest management situations, the current 
most effective methods of treatment are removal of 
heavily infected trees and isolation of infected stands 
by planting buffer zones of incompatible host species 
(Franc and Baker 2000). Even-aged silvicultural systems 
are often effective in controlling Dwarf Mistletoe, as 
the entire overstory may be removed. 

No one has quantified volume losses on individual 
Jack Pine trees due to infection by Dwarf Mistletoe. 
One study examined the effect of A. americanum on the 
growth of Lodgepole Pine (Baranyay and Safranyik 
1970). Within Manitoba, Baker et al. (1992) studied 
the impact of Dwarf Mistletoe on Jack Pine trees at the 
stand scale, focusing on loss of wood volume in the 
forest stands. The objective of this study was to com- 
pare the growth of heavily infected Jack Pine trees to 
that of non-infected or lightly infected Jack Pine trees 
based on differences in diameter at breast height (dbh), 
basal area, height, volume, and stem form. We hypoth- 
esized that there was a significant reduction in total 
cumulative dbh, basal area, height, and volume growth 
rate in trees infected with Dwarf Mistletoe. 


Methods 

The sampling area is located about 97 km north- 
east of Winnipeg, Manitoba (Figure |) within Belair 
Provincial Forest (50°38'N, 96°29'W, 250 meters 
above sea level). The underlying bedrock consists of 
Archean granites and gneisses (Manitoba Geological 
Survey 2002*). The closest meteorological station is 
located at Pine Falls about 21 km from the sampling 
area. The climate is continental (Burton et al. 1998). 
The mean annual temperature for Pine Falls is 2.1°C, 
and the total annual precipitation is 538.5 mm (Envi- 
ronment Canada 2002*). In winters, the average min- 
imum temperature reaches —23.5°C in January, and in 
summers, the average maximum temperature reaches 
25.3°C in July. 

The sampling site consists of a small, open, fire- 
originated Jack Pine stand, and is bound by Provincial 
Highway 11 to the south and a recent cutover to the 
north. The sample site is located near the south-eastern 
limit of the lodgepole pine dwarf mistletoe geograph- 
ical distribution in Manitoba (Baker et al. 1992). 
Physically, the site is characterized by level, homoge- 
neous terrain. The site is also homogeneous with regards 
to stand origin, slope variation and understory vege- 
tation. Due to the small size of the infected stand, and 
the localized nature of the mistletoe infections, the 
number of trees for sampling was limited. There are 
epicentres of Dwarf Mistletoe infections scattered 
throughout the area. Within the sampling site, there is 
a relatively even scattering of heavily infected trees 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


intermixed with lightly and non-infected trees (Fig- 
ure 1). 

Three transects were established approximately 50 m 
apart, oriented in a north-south direction, extending 
north from Provincial Hwy. 11. Ten sampling points 
were randomly established along the three transects, 
and the surrounding area was divided into four quad- 
rants. One of the four quadrants was randomly select- 
ed in which the two nearest trees to the point were select- 
ed; one heavily infected tree and the nearest lightly or 
non-infected tree between 40 and 60 years of age. 
Trees with noticeable fire scars, fungal infections, or 
injuries were avoided to minimize the effects of other 
disturbances on tree growth. Trees that exhibited exces- 
sive branching or forking on the main stem were also 
avoided to simplify stem analysis. Trees were selected 
up to 50 m from plot centre, as long as they remained 
within the stand boundaries. If no trees matching our 
criteria were found within the selected quadrant, sam- 
pling was done in the next quadrant in numerical order. 
This method helped to eliminate bias by selecting the 
closest trees to the sampling points. Once candidate 
trees were selected, they were numbered, marked with 
a north-orientation line, marked at 0.5 m and 1.3 m, and 
locations were recorded with a GPS unit (Figure 1). 
Sampling took place in late July — early August 2002. 

Prior to felling, the infestation index of the trees 
was estimated using the Hawksworth 6-class system 
(Hawksworth 1977). Infection was most easily deter- 
mined from the ground by the presence of witches’ 
brooms. Trees with an infestation index of 0 to 3 were 
classified as lightly infected, whereas trees with an 
index of 4 to 6 were classified as moderately to heavily 
infected. In order to take into account the competition 
of other trees, the dbh and distance of the nearest tree 
in each cardinal direction were measured. 

Following felling of the trees, tree height was record- 
ed and stems were sectioned at 0 m, 0.5 m, 1.3 m, and 
then 1-m intervals until stem diameter was less than 
1 cm. Sections at points of the stem where branches 
emerged were avoided, and the adjusted height was 
recorded. For each cross-section, diameter inside-bark 
and outside-bark were measured along two diameters 
perpendicular to each other. At the laboratory, age of 
each cross-section was measured, and the pointer-year 
method of cross-dating was used to validate ages (Yam- 
aguchi 1991). Following dating, stem analysis was per- 
formed on each tree to determine annual increments. 
Ring width measurements were done using the Win- 
DENDRO™ y. 2002a program (Régent Instruments 
Inc. 2002*), and annual increments were calculated us- 
ing XLSTEM™ 1.3a (Régent Instruments Inc. 1999*). 
Each cross-section was scanned with a high-resolution 
scanner and saved as a digital image. An image reso- 
lution of 800 dpi was used, except for cross-sections 
exhibiting high levels of suppression, where a resolu- 
tion of 1600 dpi was used. For one tree, suppression was 
too great for ring detection by the WinDENDRO™ 


Sast atchewan 


@ 
Winnipeg 


100° 96° g2° 


Epp AND TARDIF: EFFECS OF LODGEPOLE PINE DWARF MISTLETOE 


597] 


0 Low-infection 
O High-infection 


50 Meters 


FicurE 1: Location of the study area and distribution of the sampled trees. The lower right panel illustrates the relative posi- 
tion of the sampled trees and their infection level in the jack pine stand. Empty circles correspond to heavily infected 
trees, and empty squares correspond to lightly or non-infected trees. 


program, so measurements were performed manually 
at a magnification of 50x using a Velmex measuring 
stage to a precision of 0.001 mm. Ring widths were 
measured, starting at the pith, along four radial paths 
in the north, east, south and west directions. 

Data obtained were examined for differences between 
the two infection groups with regards to age, competi- 
tion index, dbh, basal area, height, volume and stem 
form. The competition index was modified from Hegyi’s 
distance weighted size ratio index (Avery and Burk- 
hart 2002): 

i 
Sy DIST, 


jal 

where C/I; is the competition index for the subject 
tree, D; is the dbh of the jth of four competitor trees 
(N, E, S, and W directions), D, is the dbh of the subject 
tree, and DIST, is the distance between the subject 
tree and the jth competitor tree. Basal area at dbh was 
calculated for each tree by using the formula for the 
area of a circle. Tree volume without bark was calcu- 
lated by applying Smalian’s formula for volume (Avery 
and Burkhart 2002) to each segment. The volume of 
each segment was then added to obtain tree volume. 
Stem form expressions were determined using the 
Girard form class (Avery and Burkhart 2002), with the 
quadratic mean diameter taken at the section nearest 
5 metres. Because each group had a relatively low 
sample size (N=10), the non-parametric Mann-Whitney 


U-test was performed on each variable to determine 
significant differences between samples from each 
infection group (a = 0.05). It should be noted that the 
Mann-Whitney U-test and the Wilcoxon two-samples 
test yield the same statistic and give the same results 
(Sokal and Rohlf 1997). To limit the effect of environ- 
mental variability, lightly or non-infected and heavily- 
infected trees were also carefully matched in pairs based 
on the distance separating them (Figure 1). Nine pairs 
were formed with a mean distance of 9.49 m (standard 
deviation = 4.83 m) between lightly or non-infected 
and heavily-infected trees. A Wilcoxon signed-rank 
test (Sokal and Rohlf 1997) was used to test for sig- 
nificant differences between infection groups. Paired- 
sample tests are more powerful than independent- 
sample tests. 

The cumulative dbh, basal area, height and volume 
average growth curves produced for each tree were 
compared between infection groups. First, predicted 
growth curves were modelled for each growth variable 
and tree using non-linear regression. Best fit to actual 
cumulative growth curves for each variable and tree 
were obtained using a three-parameter logistic model 
of the form: 


a 
SD 
1+ ext 
] 
where y is the value of the subject growth variable, x 
is the year of growth, a is the asymptotic level of 


598 


growth, b is the growth constant, and xp is the inflection 
point. Because each of these three parameters defines 
the shape of the growth curves, significant differences 
between parameters for the two infection groups were 
determined using the Mann-Whitney U-test (a = 0.05). 
This was repeated for each growth variable. Second, 
the best three-parameter logistic model was determined 
for each variable and infection group using the 10 trees 
as replicates. The resulting predicted growth curves 
were plotted to age 60. Because trees were not all of the 
same age and the number of observations decreased with 
time, only data covering the first 45 years of growth 
were used in all the regression analyses. 


Results 

The two infection groups showed no significant dif- 
ference in age, competition index, dbh, and basal area 
(Table 1). There was, however, a significant difference 
in height and form class (Table 1). Specifically, trees 
in the high infection group were 20.2% shorter and 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


had more stem taper. Both independent- and paired- 
sample tests yielded divergent results regarding total 
volume, and when partially controlling for environmen- 
tal variability no significant difference in total volume 
was observed as indicated by the Wilcoxon signed-rank 
test. 

The analysis of the coefficients from the logistic 
regressions derived from each tree in each infection 
group revealed that the high infection group reached a 
significantly lower maximum basal area, height, and 
volume, as predicted by the asymptotic values of the 
regression model (Table 2). Maximum dbh did not dif- 
fer significantly between groups. The logistic regres- 
sion model, projected to age 60, predicts a 21.1% lower 
basal area (Figure 2B), a 23.4% lower height (Figure 
3A), and a 42.1% lower volume (Figure 3B) for the 
high infection group. 

Results showed that there was a significantly higher 
growth rate for dbh and basal area in the high infection 
group (Table 2). This was also observed for volume but 


TABLE |: General characteristics of heavily infected and non-infected or lightly infected trees (n = 10 for each group) at time 
of sampling (Max = Maximum value, Min = Minimum value, SEM = Standard Error of the Mean). The first column of 
probabilities (P!) is based on the Mann-Whitney U-test (n=10 for each group). The second column of probabilities (P?) is 
based on the Wilcoxon signed-rank test (n=9 pairs). 


Low infection High infection 


Max Min Mean SEM Max Min Mean SEM P! Pp? 
Infestation index 2.0 0.0 12 0.29 6.0 4.0 2 0.20 0.000 0.007 
Tree age 48.0 42.0 46.3 0.62 82.0 44.0 50.0 3.59 O28 022: 
Competition index 3.34 0.77 1.85 0.30 1.89 0.66 1.29 0.15 0257) .O!515: 
Diameter at breast height (cm) 19205) W310 od 0.38 L960 SES ON eliS 45 0.83 0.544 0.515 
Basal area (m7?) 0:022> O!012. ~0:017;- 0:00 0.024 0.007 0.016 0.00 0.544 0.594 
Height (m) 14.10 10.90 12.65 0.39 L330 830 ea lORO 4s 3054: 0.004 0.008 
Volume (m?) 0.16 0.09 0.12 0.01 0.17 0.04 0.08 0.01 0.019 0.110 
Form class 0.83 0.64 0.73 0.02 0:76" 10335 0.56 0.04 0.004 0.008 


TABLE 2. Values of the three parameters of the predicted logistic growth curves used to model diameter at breast height, basal 
area, height, and volume for heavily infected and non-infected or lightly infected Jack Pine trees (n = 10 for each group). 
The three parameters are the theoretic maximum (a), growth (b), and the inflection point (x9) (Max = Maximum value, Min 
= Minimum value, SEM = Standard Error of the Mean). The first column of probabilities (P') is based on the Mann-Whit- 
ney U-test (n=10 for each group). The second column of probabilities (P?) is based on the Wilcoxon signed-rank test (n=9 
pairs). 


Low infection group High infection group 


Max Min Mean SEM Max Min Mean SEM P! pz 

Diameter at breast 
height (cm) a 22: OS na WLANs lif2,9 0.85 QO0:97 18287 2 18.97 PAIL 0.059 0.260 
b — 84) '-3230) 1 <2:46%) 10M3 -2.46 -4.29 -3.26 0.19 0.005 0.015 
Xo 36:220 23133) 28:46 1.50 Qe oie MOD3ie 524 1.19 0.023 0.028 
Basal area (m7) a 0.071 0.019 0.038 0.0061 0.032 0.007 0.016 0.0026 0.002 0.038 
b -2.32 -3.51 =O Ams ORS -3.07 -5.27 -3.84 0.20 0.001 0.008 
Xo OB Man) S464) 5538/8 Soe 39.18 24.84 31.59 1.72 0.001 0.011 
Height (m) a 24.37 14.99 19.59 1.01 20.63 9.28 13.90 1.2] 0.008 0.015 
b -1.71 -2.64 -1.98 0.09 -1.44 -2.65 -2.07 0.12 0.326 0.374 
Xo AD OI 24938 = 32°66 1.74 49.34 20.56 27.58 DATA 0.034 0.038 
Volume (m?) a 2237 2 OM2OF 1Or604s O97, 0.214 0.042 0.097 0.021 0.001 0.015 
b -2.98 -4.36 -3.60 0.13 -3.12 -6.07 -4.09 0.25 0.082 0.028 
Xo 111.79 40.48 60.48 6.51 48.74 30.68 36.59 1.97 0.001 0.011 


2004 


P<0.0001 


16 y = 16.6392/[1+(x/27.2775) 242" 
adj. ° = 0.96 
aaa, P<0.0001 eso 
5 e 
= 
a4 + 
: . y = 13,8979/{1+(x/23.3196)?°%) 
iw adj. r = 0.88 
3 P<0.0001 
5 6 
5 4 
(S) 
2 
0 
0.020 y = 0.0274/[1+(x/45.0484)22743) 
B) adj. r = 
j. F = 0.92 
P <0.0001 Wa 
~ 0.015 Vy , 
nN | i os 
© Pe 
® | | 
© 0.010 gf 
g q 
© # 
jaa) ¥ y = 0.0170/[1+(x/33.1868) 74767] 
0.005 # SAE ey 


0 10 20 30 40 50 60 
Tree age 


FIGURE 2: Average cumulative dbh growth (A) and average 
cumulative basal area growth (B) for each of the two infec- 
tion groups. Dashed lines represent predicted growth up to 
year 60 for the infection groups, based on the 3-parameter 
logistic regression models depicted. Vertical bars represent 
standard errors of the mean. The black dashed line and 
empty circles indicates the low infection group, and the 
dark grey dashed line and empty triangles indicates the 
high infection group. 


only after the Wilcoxon signed-rank test. It should be 
noted that in a logistic regression, there is an inverse 
relationship between the growth constant and the actual 
growth rate. The significantly lower growth constant 
observed in the high infection group indicates a faster 
rate of growth. No significant difference in the growth 
rate for height was observed. However, all four cumu- 
lative growth variables of the high infection group 
reached their inflection point significantly sooner than 
the low infection group (Table 2), suggesting that the 
high infection group reached its maximum growth 
rate sooner. Dbh (Figure 2A) and basal area (Figure 
2B) of the two infection groups were not predicted to 
diverge until after 45 years of age, while height growth 
curves began to diverge at about 20 years of age (Fig- 
ure 3A), and volume growth curves began to diverge 
at 35 to 40 years of age (Figure 3B). 

Results indicate a higher level of variability in the 
basal area and volume of the high-infection group when 
compared to the low-infection group, as indicated by 
the lower value of the adjusted 1 for each logistic regres- 
sion model. This may indicate variation in environmen- 
tal conditions at early stages of growth for the trees, or 


Epp AND TARDIF: EFFECS OF LODGEPOLE PINE DWARF MISTLETOE 


599 
16 y = 18.4039/(1+(x/30.4334) 1%") 
14 A) a0 f 087 iS , 
12 
E en: 
‘er Be antl 
5 : agp 
a 6 fi Fy = 13.2629/(1+(x/25.9869)29"% 
oH r=0.92 
il Bsc 
4 os 
24 a 
0 es geet 
0.16 
] B) y = 0.2048/[1+(x/46.9484)? 7] 
0.14 | adh O82 s 
0.12 i 
“E 0.10 x 
2 0.08 4 
§ ee 
S 0.06 
0.04 ft 
0.02 sf Pi y = 0.101 1/[1+(x/38.4295)?7""4 
0.00 soon? PE OOOR | 
0 10 20 30 40 me 2 
Tree age 


FIGURE 3: Average cumulative height growth (A) and average 
cumulative volume growth (B) for each of the two infection 
groups. Dashed lines represent predicted growth up to year 
60 for the infection groups, based on the 3-parameter logis- 
tic regression models depicted. Vertical bars represent stan- 
dard errors of the mean. The black dashed line and empty 
circles indicates the low infection group, and the dark grey 
dashed line and empty triangles indicates the high infec- 
tion group. 


variation in genetic resistance to mistletoe infection. 
Examination of the stem profiles also revealed that some 
trees in the high-infection group exhibited little or no 
evidence of growth reduction despite a high infection 
index (not presented). These trees could have been 
recently infected by Dwarf Mistletoe. 


Discussion 

No significant differences in tree age or competition 
levels between the two infections groups were observed 
indicating that Dwarf Misteltoe was was the key factor 
affecting Jack Pine growth. The lack of a significant 
difference in dbh between the heavily infected and 
lightly infected Jack Pine trees at the time of sampling 
likely reflects the young age of the subject trees and 
the duration of the infection period. Tinnin et al. (1999) 
found that there was a significant reduction in the 
diameter growth of Douglas-fir (Pseudotsuga menziesii 
(Mirb.) Franco var. menziesii) heavily infected by A. 
douglasii (infection index = 5 and 6) when compared 
to non-infected trees (infection index = 0) for trees hav- 
ing mean ages between 78 and 84 years. In Lodgepole 
Pine, diameter growth was not significantly affected 


600 


at stump height, but significant decreases in diameter 
growth were apparent at heights further up the stem 
because of greater stem taper (Baranyay and Safranyik 
1970). Our study also showed that stem taper was sig- 
nificantly more pronounced in the high infection group. 

At the time of sampling, basal area in the high infec- 
tion group was not significantly different than that of 
the low infection group. Comparison of the logistic 
regression coefficients provided similar results to dbh, 
except that the high infection group was predicted to 
reach a significantly lower basal area compared to the 
low infection group as trees continue to grow after 
age 45. Assuming that basal area increment after the 
age of 45 years will continue to follow that predicted by 
the logistic regression model, the projected maximum 
basal area of the high infection group was approxi- 
mately 57.9% lower than that of the low infection 
group. Pierce (1960) also found a 68.5% reduction in 
basal area for heavily infected Douglas-fir, and a 
41.0% reduction in moderately infected Douglas-fir. 

Of the significantly affected growth variables, height 
increment showed a reduction first, i.e., between the 
ages of 20 and 25 years. Pierce (1960) also found that 
differences in height were statistically significant be- 
tween all infection classes, in contrast to diameter. This 
corresponds with the highly significant 20.2% lower 
tree height in heavily infected trees in this study. The 
greater sensitivity of this variable is likely due to the 
initial response of trees to Dwarf Mistletoe infections. 
Dwarf Mistletoe infected trees accumulate a high level 
of biomass in their brooms, which detracts the allo- 
cation of resources from biomass production elsewhere 
in the tree (Tinnin and Knutson 1980). In contrast to 
the other growth variables, height of the high-infection 
group had similar variability to the low-infection group, 
as indicated by similar adjusted r values in the logistic 
regression curves. This suggests that height growth is 
less sensitive to temporal changes in factors such as 
stand density and competition (Avery and Burkhart 
2002). Height growth diverged between the two infec- 
tion groups at a much earlier age than volume, diame- 
ter, or basal area, further suggesting that height growth 
begins to show significant decreases shortly after the 
development of brooms and loss of apical dominance 
(Tinnin and Knutson 1980). 

At the time of sampling, there was a 33.3% lower 
average total volume in the high infection group, which 
was mostly due to a significant reduction in tree height. 
However, despite early reduction in height growth, 
volume growth of the heavily infected trees was not 
significantly affected until 30 to 40 years of age. Our 
values very closely correspond with those reported by 
Baranyay and Safranyik (1970), where a group of 
infected Lodgepole Pine trees with an average age of 
37 showed a 35.5% reduction in volume. From the 
logistic growth model, it was found that the predicted 
maximum average volume of the high infection group 
was 83.9% lower than the low infection group. This 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


projected value exceeded the estimated volume reduc- 
tion in infected stands made by Baker et al. (1992), 
which was between 53.4% and 70.3% depending on 
the level of potential crown closure. However, it should 
be noted that our findings reflected the volume loss in 
individual trees, and not the volume loss as averaged 
through an entire stand. 

In conclusion, this study confirmed the significance 
of the impact of Dwarf Mistletoe on the growth of com- 
mercial Jack Pine forests, and the potential economic 
loss due to severe infection. Despite early reduction in 
height growth, volume growth of the heavily infected 
trees was not significantly affected until 30 to 40 years 
of age. Further study on the effects of Dwarf Mistletoe 
over a wider range of age classes and site indexes would 
be beneficial in modelling the impact of the parasite. 


Acknowledgements 

This project was part of an undergraduate honours 
thesis course at the University of Winnipeg, conducted 
by the first author. We thank the following people for 
their contributions: F. Conciatori, G. Sayer, R. Moodie, 
R. Westwood, D. Ko Heinrichs, the University of Win- 
nipeg Department of Biology, and the Centre for For- 
est Interdisciplinary Research. Special thanks go to K. 
Knowles from Manitoba Conservation and R. Staniforth 
for their comments on a previous draft of this paper. 


Documents Cited (marked * in text) 

Environment Canada. 2002. Canadian climate normals 
1971-2000. 3 March 2003.<http://www.msc.ec.gc.ca/ 
climate/climate_normals/results_e.cfm> 

Manitoba Geological Survey. 2002. Manitoba geology. Mani- 
toba Industry, Trade and Mines. 27 March 2003. <http:// 
www.gov.mb.ca/itm/mrd/geo/exp-sup/mbgeology.html> 

Régent Instruments Inc. 1999. XLSTEM™ 1.3a. Québec, 
Quebec. 

Régent Instruments Inc. 2002. WinDENDRO™ vy. 2002a. 
Québec, Quebec. 


Literature Cited 

Avery, T. E., and H. E. Burkhart. 2002. Forest measure- 
ments. Fifth edition. Mcgraw-Hill. New York. 456 pages. 

Baker, F. A., M. Slivitsky, and K. Knowles. 1992. Impact of 
dwarf mistletoe on jack pine forests in Manitoba. Plant 
Disease 76: 1256-1259. 

Baranyay, J. A., and L. Safranyik. 1970. Effect of dwarf 
mistletoe on growth and mortality of lodgepole pine in 
Alberta. Canadian Forestry Service. Department of Fish- 
eries and Forestry. Publication Number 1285. 

Brandt, J. P., R. D. Brett, K. R. Knowles, and A. Sproule. 
1998. Distribution of severe dwarf mistletoe damage in 
west-central Canada. Natural Resources Canada, Canadian 
Forestry Service, Northern Forestry Centre, Edmonton, 
Alberta. Special Report 13. 

Burton, V. B., D. R. Zak, S. R. Denton, and S. H. Spurr. 
1998. Forest ecology, Fourth edition. John Wiley & Sons 
Inc. New York. 774 pages. 

Franc, G. D., and FE. A. Baker. 2000. Dwarf mistletoes affect- 
ing pines and firs of Wyoming. University of Wyoming and 
Utah State University. Cooperative Extension Service. B- 
1083. 


2004 


Hawksworth, F. G. 1977. The 6-class dwarf mistletoe rating 
system. USDA Forest Service, General Technical Report 
RM-48. 

Hawksworth, F. G., and D. Wiens. 1996. Dwarf mistletoes: 
biology, pathology, and systematics. Agricultural Handbook 
Number 709. USDA Forest Service. 

Hull, R. J., and O. A. Leonard. 1964a. Physiological aspects 
of parasitism in mistletoes (Arceuthobium and Phoraden- 
dron). 1. The carbohydrate nutrition of mistletoe. Plant 
Physiology 39: 996-1007. 

Hull, R. J., and O. A. Leonard. 1964b. Physiological aspects 
of parasitism in mistletoes (Arceuthobium and Phoraden- 
dron). Ul. The photosynthetic capacity of mistletoe. Plant 
Physiology 39: 1008-1017. 

Pierce, W. R. 1960. Dwarf mistletoe and its effect upon the 
larch and Douglas-fir of western Montana. Montana State 
University. School of Forestry. Missoula, Montana. Bul- 
letin Number 10. 37 pages. 

Piirto, D. D., D. L. Crews, and H. E. Troxell. 1974. The 
effects of dwarf mistletoe on the wood properties of lodge- 
pole pine. Wood and Fiber 6: 26-35. 

Rudolph, T. D., and P. R. Laidly. 1990. Pinus banksiana Jack 
Pine. Jn Silvics of North America: |. Conifers; 2. Hard- 
woods. Agriculture Handbook 654. U.S. Department of 
Agriculture, Forest Service, Washington, DC. Volume 2. 
Technical coordinators, Burns, R. M., and B. H. Honkala. 
877 pages. 


Epp AND TARDIF: EFFECS OF LODGEPOLE PINE DWARF MISTLETOE 60) 


Sala, A., E. V. Carey, and R. M. Callaway. 2001. Dwarf 
mistletoe affects whole-tree water relations of Douglas 
fir and western larch primarily through changes in leaf to 
sapwood ratios. Oecologia 126: 42-52. 

Sims, R. A., H. M. Kershaw, and G. M. Wickware. 1990. 
The autecology of major tree species in the north central 
region of Ontario. Technical Report 48, Northwestern 
Ontario Forest Technology Development Unit, Ontario 
Ministry of Natural Resources, Thunder Bay, Ontario. 
126 pages. 

Sokal, R. R., and F. J. Rohlf. 1997. Biometry. W.H. Free- 
man and Company, New York. 

Tinnin, R. O., and D. M. Knutson. 1980. Growth charac- 
teristics of the brooms on Douglas-fir caused by Arceutho- 
bium douglasii. Forest Science 26: 149-158. 

Tinnin, R. O., C. G. Parks, and D. M. Knutson. 1999. 
Effects of Douglas-fir dwarf mistletoe on trees in thinned 
stands in the pacific northwest. Forest Science 45(3): 
359-365. 

Yamaguchi, D. K. 1991. A simple method for cross-dating 
increment cores from living trees. Canadian Journal of 
Forest Research 21: 414-416. 


Received 10 February 2004 
Accepted 24 December 2004 


Notes 


Excavation of an Arctic Fox, Alopex lagopus, den by a Polar Bear, 


Ursus maritimus 


Evan S. RICHARDSON! and RYAN K. BROOK? 


‘Canadian Wildlife Service, 5320-122 St., Edmonton, Alberta T6H 3S5 Canada 
Faculty of Environment, University of Manitoba, Winnipeg, Manitoba R3T 2N2 Canada 


Richardson, Evan S., and Ryan K. Brook. 2004. Excavation of an Arctic Fox, Alopex lagopus, den by a Polar Bear, Ursus 


maritimus. Canadian Field-Naturalist 118(4): 602-603. 


We observed a Polar Bear (Ursus maritimus) excavating an Arctic Fox (Alopex lagopus) den on 24 June 1998, 3 km inland from 


the Hudson Bay coast (58°40'N, 93°12'W), near Cape 
Churchill in Wapusk National Park, Manitoba. To our 
knowledge this is the first observed excavation of an Arctic 
Fox den by a Polar Bear. 


Key Words: Arctic Fox, Alopex lagopus, Polar Bear, Ursus 
maritimus, den, behaviour, Hudson Bay, Wapusk 
National Park, Manitoba. 

The western Hudson Bay Polar Bear (Ursus mar- 
itimus) population remains on shore from approximate- 
ly mid-July through early November due to complete 
annual melting of the sea ice (Stirling et al. 1977). 
During this period, bears mainly remain relatively 
inactive (Knudsen 1978; Latour 1981; Lunn and Stir- 
ling 1985) and subsist primarily on stored fat reserves 
(Nelson et al. 1983; Derocher et al. 1990; Ramsay et 
al. 1991). However, during the summer months, polar 
bears are opportunistic and occasionally feed on a 
variety of terrestrial food sources including grasses, 
sedges and berries (Knudsen 1978; Lunn and Stirling 
1985, Derocher et al. 1993), Thick-billed Murres (Uria 
lomvia) (Donaldson et al. 1995), Canada Geese (Bran- 
ta canadensis) (Russell 1975) and their eggs (Smith 
and Hill 1996), Snow Goose eggs (Anser caerulescens) 
(Abraham et al. 1977), Willow Ptarmigan (Lagopus 
lagopus) (Miller and Woolridge 1983), seabirds (Stemp- 
niewicz 1993), microtine rodents (Russell 1975; Miller 
and Woolridge 1983), and Caribou (Rangifer tarandus) 
remains (Brook and Richardson 2002). 

We observed an adult male Polar Bear excavate an 
Arctic Fox den near Nestor | field camp (58°40'N, 
93°12'W), near Cape Churchill in Wapusk National 
Park, Manitoba on 24 June 1998 at 11:00 CST. The 
bear was initially sighted on a large beach ridge inves- 
tigating one of the camp buildings and then moved to 
an Arctic Fox den located approximately 350 m south- 
east of camp. The bear moved around the site, inves- 
tigated several of the den entrances and then stopped 
at one entrance and began to excavate the den. The 
bear stopped digging every 10 —15 seconds to put its 


head into the enlarged entrance, continued this behav- 
iour for approximately 3 minutes, then stopped dig- 
ging and spent approximately 2 minutes investigating 
several other den entrances, but did not dig them out. 
The bear then moved away from the den and down 
the eastern side of the beach ridge, where it could no 
longer be observed. The bear was not observed eating 
anything, although while its head was in the entrance, 
it could not be observed. Several fox pups were occu- 
pying the den at the time and we assumed the pups 
were in the den during the observation although the 
location of the vixen at the time was unknown. We 
visited the following day and found a large pit dug by 
the bear. The excavation was 55 cm at its deepest and 
was approximately 70 cm wide and 80 cm in length. 
Arctic Fox pups could still be heard inside the den 
confirming that the den had not been abandoned and 
there was no evidence any fox pups were killed. The 
top of the den was littered with Canada and Snow 
Goose remains, consisting mostly of feet (98) and other 
waterfowl remains such as bones and feathers. Cached 
prey items inside the den were visible from several den 
entrances and consisted primarily of goose remains. 
Food remains found at fox dens in the region included 
both adult and juvenile geese, ducks, Caribou remains 
and in one instance, a Muskrat (Ondatra zibethicus) 
(Richardson and Brook, personal observations, 1999). 

Polar Bears are known to excavate subnivean lairs 
in search of young Ringed Seal (Phoca hispida) pups 
(Stirling and Archibald 1977). They also excavate earth 
and snow dens for reproduction, as well as open pits 
for resting (Clark 1996; Clark et al. 1997). Although 
adult male bears are known to excavate open pits on 
coastal beach ridges (Clark 1996), we suggest that the 
bear was most probably trying to gain access to a poten- 
tial food source. The bear may have been attracted to 
the fox den for several reasons. In areas where prey is 
abundant, Arctic Foxes cache large quantities of food 
at den sites for later consumption (Sklepkovych and 


602 


2004 


Montevecchi 1996; Garrott et al. 1984). As a result, 
other predators may be attracted to den sites by their 
smell (Prestrud 1992). At active den sites, character- 
| istic barks of arctic fox pups can be heard from with- 
_ in the den when it is disturbed (Eberhardt et al. 1983; 
Richardson and Brook, personal observations, 1999). 
Bears are adept at locating food sources by smell (Lunn 
and Stirling 1985) and it seems unlikely that barking 
attracted the bear to the site, but the sound may have 
stimulated further investigation. Prey remains at den 
_ sites may provide a direct energy source for bears, how- 
| ever the energetic cost of excavating a den would not 
likely be repaid in the capture of a small Arctic Fox pup. 
Although, Macpherson (1969) notes that Arctic Fox 
dens may be occasionally excavated by Grizzly Bears 
| (Ursus arctos) and Wolves (Canis lupus), we are not 
aware of any other published reports of Polar Bears 
| excavating Arctic Fox dens. 


| Acknowledgments 

These observations were made while conducting 
research supported by Wapusk National Park, the 
Manitoba Department of Conservation, the Churchill 
_ Northern Studies Centre, the Northern Studies Training 
_ Program of the Department of Indian and Northern 
Affairs Canada, the Canadian Department of Fisheries 
and Oceans, the Western Canada Service Centre of 
_ Parks Canada, Wat’chee Lodge, the Canadian Wildlife 
Federation, and the Manitoba Chapter of the Wildlife 
Society. We would like to thank I. Stirling and N. J. 
Lunn for their valuable comments on the manuscript. 


Literature Cited 

Abraham, K. F., P. Mineau, and F. Cooke. 1977. Unusual 
predators of snow goose eggs. Canadian Field-Naturalist 
91: 317-318. 

Brook, R. K., and E. S. Richardson. 2002. Observations of 
polar bear, Ursus maritimus, predatory behavior toward 
caribou, Rangifer tarandus, in Wapusk National Park, 
Manitoba. Arctic 55: 193-196. 

Clark, D. A. 1996. Terrestrial habitat selection by Polar Bears 
(Ursus maritimus Phipps) in the Western Hudson Bay 
Lowlands. M.Sc. thesis, University of Alberta, Edmonton, 
Alberta. 

Clark, D. A., I. Stirling, and W. Calvert. 1997. Distribution, 
characteristics and use of earth dens and related excavations 
by polar bears on the western Hudson Bay lowlands. Arctic 
50: 158-166. 

Derocher, A. E., R. A. Nelson, I. Stirling, and M. A. Ram- 
say. 1990. Effects of fasting and feeding on serum urea and 
serum creatine levels in polar bears. Marine Mammal 
Science 6: 196-203. 

Derocher, A. E., D. Andriashek, and I. Stirling. 1993. Ter- 
restrial foraging by polar bears during the ice-free period 
in western Hudson Bay. Arctic 46: 251-254. 

Donaldson, G. M., G. Chapdelaine, and J. D. Andrews. 
1995S. Predation of thick-billed murres, Uria lomvia, at two 


NOTES 


603 


breeding colonies by polar bears, Ursus maritimus, and 
walruses, Odobenus rosmarus. Canadian Field-Naturalist 
109: 112-114. 

Eberhardt, L. E., R. A. Garrott, and W. C. Hanson. 1983. 
Den use by arctic foxes in northern Alaska. Journal of 
Mammalogy 64: 97-102. 

Garrott, R. A., L. E. Ebehardt, and W. C. Hanson. 1984. 
Arctic fox denning behaviour in northern Alaska. Cana- 
dian Journal of Zoology 62: 1636-1640. 

Knudsen, B. 1978. Time budgets of polar bears, Ursus mar- 
itimus, on North Twin Island, James Bay, during summer. 
Canadian Journal of Zoology 56: 1627-1628. 

Latour, P. B. 1981. Spatial relationships and behaviour of 
polar bears (Ursus maritimus Phipps) concentrated on land 
during the ice-free season of Hudson Bay. Canadian Jour- 
nal of Zoology 59: 1763-1774. 

Lunn, N. J., and I. Stirling. 1985. The significance of sup- 
plemental food to polar bears during the ice-free period 
of Hudson Bay. Canadian Journal of Zoology 63: 2291- 
2297. 

Macpherson, A. H. 1969. The dynamics of Canadian arctic 

fox populations. Canadian Wildlife Service Report Series 

8. 52 pages. 

Miller, G. D., and D. R. Woolridge. 1983. Small game hunt- 

ing behaviour of polar bears, Ursus maritumus. Canadian 

Field-Naturalist 97: 93-94. 

Nelson, R. A., G. E. Folk Jr., E. W. Pfeiffer, J. J. Craig- 
head, C. J. Jonkel, and D. L. Steiger. 1983. Behavior, 
biochemistry, and hibernation in black, grizzly, and polar 
bears. International Conference on Bear Research and 
Management 5: 284-290. 

Prestrud, P. 1992. Denning and home-range characteristics 
of breeding arctic foxes in Svalbard. Canadian Journal of 
Zoology 70: 1276-1283. 

Ramsay, M. A., R. A. Nelson, and I. Stirling. 1991. Season- 
al changes in the ration of serum urea to creatinine in feed- 
ing and fasting polar bears. Canadian Journal of Zoology 
69: 298-302. 

Russell, R. H. 1975. The food habits of polar bears of James 
Bay and southwest Hudson Bay in summer and autumn. 
Arctic 28: 117-129. 

Sklepkovych, B. O., and W. A. Montevecchi. 1996. Food 
availability and food hoarding behaviour by red and arc- 
tic foxes. Arctic 49: 228-234. 

Smith, A. E., and M. R. J. Hill. 1996. Polar bear, Ursus mar- 
itimus, depredation of Canada goose, Branta canadensis, 
nests. Canadian Field-Naturalist 110: 339-340. 

Stempniewicz, L. 1993. The Polar bear, Ursus maritimus, 
feeding in a seabird colony in Frans Josef Land. Polar 
Research 12: 33-36. 

Stirling, I., and W. R. Archibald. 1977. Aspects of predation 
of seals by polar bears. Journal of the Fisheries Research 
Board of Canada 34: 1126-1129. 

Stirling, I., C. Jonkel, P. Smith, R. Robertson, and D. 
Cross. 1977. The ecology of the polar bear (Ursus mar- 
itimus) along the western coast of Hudson Bay. Canadian 
Wildlife Service Occasional Paper 33: 64 pages. 


Received 24 June 2003 
Accepted 2 November 2004 


604 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Two Great Black-backed Gulls, Larus marinus, Kill Male Longtailed 


Duck, Clangula hyemalis 


MARTHA DowsLey! and ANDREW CIRTWILL? 


‘Department of Geography, McGill University, 805 Sherbrooke Street West, Montreal, Quebec H3A 2K6 Canada 


°515 Roosevelt Drive, Kingston, Ontario K7M 5Y2 Canada 


Dowsley, Martha, and Andrew Cirtwill. 2004. Two Great Black-backed Gulls, Larus marinus, Kill Male Longtailed Duck, 
Clangula hyemalis. Canadian Field-Naturalist 118(4): 604-605. 


At Presqu’ile Point, Presqu’ile Provincial Park, Ontario on 23 March 2003, while interpreting the waterfowl migration for 
park visitors, we witnessed two adult Great Black-backed Gulls attack and kill a male Long-tailed Duck. 


Key Words: Great Black-backed Gull, Larus marinus, Long-tailed Duck, Clangula hyemalis, predation 


Great Black-backed Gulls, Larus marinus, are known 
to be scavengers and to take eggs, nestlings, and fledg- 
lings of other birds. However, their efficacy as preda- 
tors is less well documented. On 23 March 2003 we 
witnessed two adult Great Black-backed Gulls attack 
and kill a Long-tailed Duck, Clangula hyemalis. 

We were using a spotting scope (power 45x) to 
observe diving ducks located 200 to 250 m from shore 
off Presqu’ile Point, Ontario (44°00'N, 77°41'W) as they 
fed and rested at the front edge of ice in Presqu’ile 
Bay. The ducks routinely lifted off when a predatory 
bird passed overhead. After one such take-off, we ob- 
served a pair of adult Great Black-backed Gulls flying 
above the ducks. One male Long-tailed Duck failed 
to lift off and the gulls flew down and began to harass it. 
We did not see the initial separation of the duck from 
the flock, so we were unable to observe whether it was 
healthy and merely caught under water when the rest of 
the flock flew, or whether it was already injured when 
attacked. 

The attack proceeded in what appeared to be a coor- 
dinated effort by the gulls. One Great Black-backed 
Gull flew in circles overhead when the duck dived and 
swam under water. When the duck surfaced, the gull 
alighted on the water and tried to bite him. The sec- 
ond gull then took off from the water and flew over- 
head, watching the duck as he dived again to avoid 
the attack of the first gull. When the duck resurfaced, 
the second gull attacked him while the first gull again 
took off and watched as the duck tried to make another 
escape under water. 

The Great Black-backed Gulls continued to take 
turns flying and attacking the duck for 20-25 minutes. 
At the end of this time, one gull succeeded in grabbing 
the wing of the Long-tailed Duck and both gulls then 
pecked at the body of the duck. Less than 5 minutes 
later the duck was dead, and its limp body could be seen 
floating on the water. The adult Great Black-backed 
Gulls were then joined by an immature Great Black- 
backed Gull and all three gulls began to eat the dead 
duck. We could not continue observations as the wind 
and currents had pushed the duck’s carcass out into Lake 
Ontario, beyond the useful range of our spotting scope. 


Great Black-backed Gulls have a varied diet, but their 
main food sources are fish and seabirds (both often 
consumed as carrion) (Buckley 1990). Their predation 
of birds is focused on the eggs, nestlings, newly fledged 
young and sick or injured individuals (Beaman 1978; 
Mawhinney and Diamond 1999). Healthy adult birds 
are rarely killed, with the notable exception of preda- 
tion in some seabird colonies, such as those of the 
Atlantic Puffin (Fratercula arctica) in Northern Europe 
(Beaman 1978), and Newfoundland (Nettleship 1972). 

Great Black-backed Gulls are considered to be pred- 
ators of waterfowl (Good 1998); however, observations 
of Great Black-backed Gulls attacking and killing adult 
ducks are quite rare. One notable record reported by 
Cleghorn (1942) was of an adult Great Black-backed 
Gull killing a female Goldeneye (Bucephala clangula) 
and an immature Great Black-backed Gull killing anoth- 
er unidentified duck the same day. 

Cobb (1957) observed a single adult Great Black- 
backed Gull kill an injured Ruddy Duck, Oxyura 
Jamaicensis, in Rhode Island. In that case the method 
of attack was similar to what we witnessed at Presqu ile. 
The gull hovered above the duck as it dived and then 
attempted to grab it when it resurfaced (Cobb 1957). 
Our report corroborates the evidence that Great Black- 
backed Gulls will fly above diving ducks, presumably 
to watch them under water, and attack them when they 
resurface. 

Addy (1945) witnessed a group of Great Black- 
backed Gulls attack and kill an adult American Black 
Duck, Anas rubripes. The gulls chased and swooped 
at groups of American Black Ducks until one adult 
gull injured a duck. The gulls then worried the duck by 
lifting it from the water by its neck and by grasping 
its back. After about half an hour the duck died, and 
was consumed, mainly by the gull which had first 
attacked it (Addy 1945). This observation does not 
appear to have been a coordinated attack on the duck 
as in the case we witnessed, because one gull of a 
group carried out most of the attack and was the pri- 
mary consumer of the carcass. 

In our observation, the two Great Black-backed Gulls 
showed a level of coordination in their attack that is 


On 


2004 


unrecorded in the literature. The method they used of 
taking turns flying overhead and attacking the duck 
seems to be an efficient means to harass their prey to 


| the point of exhaustion and death. 


Literature Cited 

Addy, C. E. 1945. Great Black-backed Gull kills adult Black 
Duck. Auk 62: 142-143. 

Beaman, M. A. S. 1978. The feeding and population ecology 
of the Great Black-backed Gull in northern Scotland. Ibis 
120: 126-127. 

Buckley, N. J. 1990. Diet and feeding ecology of Great 
Black-backed Gulls (Larus marinus) at a southern Irish 
breeding colony. Journal of Zoology: Proceedings of the 
Zoological Society of London 222: 363-373. 


GLYNNIS A. Hoop and TIM NEUFELD 


NOTES 


605 


Cleghorn, J. D. 1942. Great Black-backed Gull killing Amer- 
ican Goldeneye. Auk 59: 584-585. 

Cobb, S. 1957. Great Black-backed Gull (Larus marinus). 
Auk 74: 498. 

Good, T. 1998. Great Black-backed Gull. The birds of North 
America (330). Edited by A. Poole and F. Gill in The 
Birds of North America, Inc., Philadelphia Pennsylvania. 

Mawhinney, K., and A. W. Diamond. 1999. Using radio- 
transmitters to improve estimates of gull predation on Com- 
mon Eider ducklings. Condor 101: 824-831. 

Nettleship, D. N. 1972. Breeding success of the Common 
Puffin (Fratercula arctica) on different habitats at Great 
Island, Newfoundland. Ecological Monographs 42: 239- 
268. 


Received 7 May 2004 
Accepted 4 October 2004 


| First Record of Mountain Lions, Puma concolor, in Elk Island National 
| Park, Alberta 


| Elk Island National Park, R.R. #1, Site 4, Fort Saskatchewan, Alberta T8L 2N7 Canada, e-mail: [Hood] glynnis.hood@pc.ge.ca 


| Hood, Glynnis A., and Tim Neufeld. 2004. First record of Mountain Lions, Puma concolor, in Elk Island National Park, 


Alberta. Canadian Field Naturalist 118(4): 605-607. 


Several sightings of Mountain Lions (Puma concolor) and wildlife mortalities consistent with predation by Mountain Lion have 
occurred in and adjacent to Elk Island National Park from February 2003 to present. These are the first recorded Mountain 


| Lions (locally called Cougar) sightings since the area encompassing the park was protected in 1906. 


Key Words: Mountain Lion, Cougar, Puma, Panther, Catamount, Puma concolor, Elk Island National Park, Alberta, range, 


first record. 


Along with its current range, Mountain Lion (Puma 
concolor) historically occupied central and eastern 
Canada (Banfield 1974). In present day Aiberta how- 
ever, Mountain Lion (locally called Cougar) popula- 
tions are mainly concentrated in the Rocky Mountains 
and the foothills (Soper 1964; Banfield 1974; Smith 
1993; Pattie and Fisher 1999). By 1961, Bird described 
many large predators, including the Mountain Lion, as 
being extirpated from the aspen parkland of Canada’s 
prairie provinces. Soper (1964) referenced individual 


_ reports of Mountain Lions north of Edmonton, Alberta 
_ at Boiler and Grand Rapids on the Athabasca River 


and in the Peace River Region in northwestern Alberta. 


_ He also noted occasional sightings in southern Alberta 


along the Milk, Bow, and South Saskatchewan River 
systems, as well as in the Cypress Hills. Smith (1993) 
added additional sightings near the towns of White- 
court and Athabasca, Alberta. 

Sightings of Mountain Lions are increasing in areas 
where they were previously unreported. Occasional 
sightings have been documented to the north in Wood 
Buffalo National Park (Gau et al. 2001), and to the 
south and west of Elk Island National Park (EINP) near 
Cooking Lake, Pigeon Lake, and Sherwood Park (Smith 
1979). However, until 2003, there had never been a 


documented Mountain Lion sighting in EINP (Soper 
1940*; Soper 1951; Burns and Cool 1984*; Parks 
Canada Warden Service unpublished notes). Elk Island 
lies between longitude W112°57'00" and W112°46'45" 
and latitude N53°42'57" and N53°30'37" (Figure 1) 
and covers an area of 196 km*. The park is located in 
the Aspen Parkland Natural Subregion (Achuff 1994) 
in the Beaver Hills of east-central Alberta and is home 
to several species of large ungulates. 

Given the known distances that Mountain Lions 
would have to travel to access the park, Burns and 
Cool (1984) predicted “occurrences of cougar in Elk 
Island National Park [EINP] in future remain a slim 
possibility”. The closest source populations to EINP 
are in the Rocky Mountains foothills and the Swan Hills 
region of north-central Alberta (Pattie and Fisher 1999). 
These populations are approximately 200 km away 
from EINP, which is well within the 500 km dispersal 
distance observed for Mountain Lions (Logan and 
Sweanor 1999). 

On 8 February 2003 at 11:00 h one of us observed 
an adult Mountain Lion approximately 10 km south 
of the boundary of EINP at Islet Lake in the southern 
reaches of the Cooking Lake — Blackfoot Grazing, 
Wildlife and Provincial Recreation Area (Figure 2). 


606 


,EBERTA 


Elk Island 
National Park 


Edmonton @ 


, =» Calgary == Mountain lion 


N distribution 
A in Alberta 
6 i 10 = w Es 


FIGURE |. Location of Elk Island National Park in east-central 
Alberta. The shaded area indicates the distribution of 
Mountain Lion in Alberta (based on Pattie and Fisher 
1999). 


In late March, a conservation officer found tracks of 
one large Mountain Lion and a much smaller Moun- 
tain Lion together near the northern boundary of the 
Provincial Recreation Area — an area that bounds the 
southern extent of EINP. On 26 July 2003 at 14:00 h 
a group of Young Canada Works students working in 
the park reported seeing an adult Mountain Lion just 
south of Highway 16 in the southern half of EINP. 

An adult Mountain Lion was also seen by park vis- 
itors on two different occasions in the northcentral 
section of the park on 29 July 2003. The first visitor 
reported seeing a Mountain Lion at 07:30 h beside 
the main road through the park and a second visitor 
saw a Mountain Lion while on a nearby trail at 21:40 h. 
In the latter case the visitor reported seeing the Moun- 
tain Lion watching a herd of Bison (Bison bison) 
approximately 5 km from the trailhead. Equestrians 
saw two Mountain Lions together in the north central 
part of the Provincial Recreation Area around the same 
time. 

Further sightings were reported in the fall of 2003 
when a park visitor observed an adult Mountain Lion 
while hiking on 17 September. Another park visitor 
reported seeing a Mountain Lion cross the main park 
road on 2 October. In mid-October, a hiker reported 
seeing a Mountain Lion at the southern part of the 
Provincial Recreation Area between Push Lake and 
Islet Lake. On 20 November 2003 a park warden report- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Elk Island N 


nal.Park 


‘{ » 
atio 
iN 


(i 


Sighting 
Ungulate Mortality 


CookingLake - Blackfoot Grazing, Wildlife 
and Provincial Recreation Area 


FiGure 2. Mountain Lion sightings and wildlife mortalities 
consistent with predation by Mountain Lion in Elk 
Island National Park and the Cooking Lake — Black- 
foot Grazing, Wildlife and Provincial Recreation Area 
since February 2003. 


ed seeing a Mountain Lion crossing a secondary road 
in the east-central part of EINP. 

In 2004, a cyclist observed a Mountain Lion sitting 
beside the main road in the northern part of the park 
(13 July). The most recent confirmed sighting in the 
park was on 24 October 2004 when a park resident 
observed a Mountain Lion in the main part of the park 
(Figure 2). On two occasions, acreage owners on lands 
east and north of the park heard estrous calls of a Moun- 
tain Lion (October 2003 and August 2004). The land- 
owner who reported the August 2004 calls observed a 
small Mountain Lion on his property a few days later. 

In addition to sightings, four ungulate mortalities 
with wounds consistent with Mountain Lion predation 
(Ross et al. 1997) have been documented in the park 
since the winter of 2002-2003 (Figure 2). Prey species 
were: 2 American Elk (Cervus canadensis), 1 Moose 
(Alces alces), and 1 White-tailed Deer (Odocoileus 
virginianus). In all cases the carcasses had signs of 
wounds and fatal injuries to the neck and, in some 
cases, large puncture marks in the hide. Apart from 
Coyotes (Canis latrans), EINP lacks any large resident 
predators. In addition, the park has very high ungulate 
densities (approximately 13 ungulates per km7; Parks 
Canada unpublished data) and is home to American 
Elk, Moose, Bison (Bison bison), White-tailed Deer, 
and Mule Deer (Odocoileus hemionus). The park also 
has a large Beaver (Castor canadensis) population and 
many resident medium and small mammal species 


| including porcupine (Erethizon dorsatum), Muskrat 
| (Ondatra zibethicus) and Red Squirrel (Jamiasciurus 
| hudsonicus). Hare (Lepus spp.) are at low densities in 


the park (Super 1951). 

The park provides adequate undeveloped habitat and 
food resources to support an adult Mountain Lion 
(Spalding and Lesowski 1971; Ross and Jalkotzy 1992; 
Ross et al. 1997; Robinson et al. 2002). Currently, park 
staff translocate Bison and American Elk away from 
the park to reduce the impact of intense herbivory on 
vegetation. The presence of large predators, such as 
Mountain Lions, may help to reduce the ungulate den- 


| sities in the park, and therefore moderate the amount of 
| active management of ungulate populations by park 
| staff. 


The park is completely fenced and the abundance of 
prey species might offer enough resources to support a 


| future Mountain Lion population. Given that success- 

| ful reproduction in Mountain Lions is related to the 

| availability of food (Pierce et al. 2000), the likely pres- 

'ence of a mated pair within EINP suggests such a 
scenario 1s not unrealistic. 


Acknowledgments 
We thank Conservation Officer Ed Whitelock and 
Park Warden Olaf Jensen for providing us with addi- 


( tional information on Mountain Lion sightings. We 


also thank three anonymous reviewers for their edito- 
rial comments. 


Documents Cited 


| Burns, G. R., and N. L. Cool. 1984. A biophysical inventory 


of the non-ungulate mammals of Elk Island National Park. 
Report prepared for Parks Canada by the Canadian Wild- 
life Service, Edmonton, Alberta, Canada. 195 pages. 

Soper, J. D. 1940. Preliminary final report on Elk Island Na- 
tional Park, Alberta, Canada. unpublished report. Depart- 
ment of Mines and Resources Lands, Parks and Forest 
Branch, National Parks Bureau. Edmonton, Alberta, Cana- 
da. 46 pages. 


Literature Cited 

Achuff, P. L. 1994. Natural regions, subregions and natural 
history themes of Alberta — a classification for protected 
areas management, revised and updated December 1994. 


NOTES 


607 


Alberta Environmental Protection, Edmonton, Alberta, 
Canada. 72 pages. 

Banfield, A. W. F. 1974. The mammals of Canada. Univer- 
sity of Toronto Press, Toronto, Ontario, Canada. 438 pages. 

Bird, R. D. 1961. Ecology of the aspen parkland of western 
Canada in relation to land use. Canada Department of Agri- 
culture, Ottawa, Ontario, Canada. 155 pages. 

Gau, R. J., R. Mulders, T. Lamb, and L. Gunn. 2001. 
Cougars (Puma concolor) in Northwest Territories and 
Wood Buffalo National Park. Arctic 54: 185-187. 

Logan, K. A., and L. Sweanor. 1999. Puma. Pages 347-77 
in Ecology and management of large mammals in North 
America. Edited by S. Demarais and P. R. Krausman 
Prentice-Hall, Englewood Cliffs, New Jersey. 

Pattie, D., and C. Fisher. 1999. Mammals of Alberta. Lone 
Pine Publishing. Edmonton, Alberta, Canada. 240 pages. 

Pierce, B. M., V. C. Bleich, and R. T. Bowyer. 2000. Social 
organization of Mountain Lions: does a land-tenure sys- 
tem regulate population size? Ecology 81: 1533-1543. 

Robinson, H. S., R. B. Wielgus, and J. C. Gwilliam. 2002. 
Cougar predation and population growth of sympatric mule 
deer and white-tailed deer. Canadian Journal of Zoology 
80: 556-568. 

Ross, P. I., and M. G. Jalkotzy. 1992. Characteristics of a 
hunted population of cougars in southwestern Alberta. 
Journal of Wildlife Management 56: 417-426. 

Ross, P. I., M. G. Jalkotzy, and B. M. Festa. 1997. Cougar 
predation on bighorn sheep in southwestern Alberta dur- 
ing winter. Canadian Journal of Zoology 75: 771-775. 

Smith, H. C. 1979. Mammals of the Edmonton area. Provin- 
cial Museum of Alberta Natural History Occasional Paper 
Number 2. Queens Printer, Edmonton, Alberta, Canada. 
34 pages. 

Smith, H. C. 1993. Alberta mammals, an atlas and guide. The 
Provincial Museum of Alberta. Edmonton, Alberta, Cana- 
da. 238 pages. 

Soper, J. D. 1951. The mammals of Elk Island National Park, 
Alberta Canada. Wildlife Management Bulletin Series 1, 
Number 3. Canadian Wildlife Service, Ottawa, Ontario, 
Canada. 27 pages. 

Soper, J. D. 1964. The mammals of Alberta. The Hamley 
Press, Ltd. Edmonton, Alberta, Canada. 402 pages. 

Spalding, D. J., and J. Lesowski. 1971. Winter food of the 
cougar in south-central British Columbia. Journal of Wild- 
life Management 35: 378-381. 


Received 19 December 2003 
Accepted 23 December 2004 


608 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


First Confirmed Occurrence of a Wolf, Canis lupus, South of the 
St. Lawrence River in Over 100 Years 


Mario VILLEMURE! and HELENE JOLICOEUR? 


'Université de Sherbrooke, Département de Biologie, Sherbrooke, Québec JIK 2R1 Canada Current address: 11A, chemin 
St-Francois, St-Mathieu-du-Parc, Québec GOX INO Canada 
*Société de la faune et des parcs du Québec, Direction du développement de la faune, 675, boulevard René-Lévesque Est. . 


11" floor, box 92, Québec, Québec GIR 5V7 Canada 


Villemure, Mario, and Helene Jolicoeur. 2004. First confirmed occurrence of a Wolf, Canis lupus, south of the St. Lawrence 
River in over 100 years. Canadian Field-Naturalist 118(4): 608-610. 


A large canid was snared near Sainte-Marguerite-de-Lingwick, Québec, in January 2002. DNA analysis confirmed the ani- 
mal to be a Wolf (Canis lupus). Wolves were extirpated from this region around 1850-1900 and this is the first confirmed 


observation since then. 


Key Words: Wolf, Canis lupus, range, dispersal, St. Lawrence River, Québec. 


Historically, Wolves (Canis lupus) were distributed 
across most of North America. They were extirpated 
in the southern portion of their range by the turn of 
the century (Nowak 1983). In Québec, Wolves disap- 
peared from the south shore of the St. Lawrence River 
around 1850-1900 (Peterson 1966). The extermination 
of the Wolf and the development of agriculture in the 
mid-1900 facilitated the extension of Coyote (Canis 
latrans) range. Wolves are now mostly limited to the 
northern and less populated regions of North America 
(Wayne et al. 1992). 

On 19 January 2002, a male Wolf was trapped 
near the village of Sainte-Marguerite-de-Lingwick 
(45°36'15"N, 71°17'15"W) in the Eastern Townships 
of southern Québec. The Wolf weighed 29.1 kg, sim- 
ilar to the weight of an adult from Papineau-Labelle 
(Potvin 1986*) or a yearling from the Laurentides 
region (Jolicoeur 1998%*). A tissue sample was collected 
from the temporal muscle for genetic identification 
of the species. DNA analyses were performed by the 
Natural Resources DNA Profiling and Forensic Center 
(Trent University, Peterborough, Ontario K9J 7B8 Can- 
ada) following the method described in Wilson et al. 
(2000). The Eastern Townships sample was profiled at 
the mitochondrial DNA (mtDNA) control region and 
8 microsatellite loci. The genetic profile was compared 
to samples of canids representing the Eastern Wolf (C. 
lupus lycaon) from Algonquin Provincial Park; West- 
ern Coyotes (C. latrans) from Texas, North Carolina 
and Ohio; and Wolves (C. lupus) from Pukaskwa Na- 
tional Park, northeastern Ontario and the Laurentide 
Wildlife Reserve region north of Québec City. The 
above populations were used to compare the Eastern 
Townships sample against C. lupus lycaon, C. latrans 
and C. lupus to assess the species-of-origin or hybrid 
genotype. 

The sample had a mtDNA consistent with C. 1. lycaon/ 
C. latrans and the microsatellite genotype suggested 
95.0% shared ancestry with Eastern Wolves from 
Algonquin Provincial Park. The sample had a DNA 
profile consistent with an Eastern Wolf. The Eastern 


Wolf is generally described as being smaller than other 
Gray Wolf subspecies (Nowak 1995, 2002). Although 
it has been proposed as a distinct species (C. lycaon) 
by Wilson et al. (2000), its status as a subspecies is 
still generally accepted. Some authors suggest it may 
result from hybridization between C. rufus and C. 
Tupus (Nowak 2002). In Québec, the Eastern Wolf is 
found mostly in the southern deciduous and mixed 
forests (Jolicoeur and Henault 2002*) Coyote genes 
found in the mitochondrial DNA of the specimen have 
been reported before in Wolves in eastern Canada 
(Lehman et al. 1991; Wilson et al. 2000). Wolves and 
Coyotes are most likely to interbreed when Wolf den- 
sity is low relative to Coyotes and when the species 
are similar in size. In southern Québec, male Coyotes 
weigh 14.6—18.7 kg (Fortin and Huot 1995*; Dumond 
and Villard 2000; Villemure 2003*). 

Wolf dispersal has been monitored in southern Qué- 
bec (Messier 1985; Potvin 1987: Jolicoeur 1998*; 
Villemure 2003*, but Wolves have not been docu- 
mented crossing the St. Lawrence River. Harrison and 
Chapin (1998) identified two potential corridors link- 
ing Wolf populations north of the St. Lawrence River 
to potential habitat in Maine and New Hampshire. 
However, movements of Wolves south of their current 
range are thought to be unlikely because of potential 
barriers such as the St. Lawrence Seaway and regions 
with high human population, high road density and 
intensive agriculture (Wydeven et al. 1998). Potential 
core habitat for Wolves has been identified in New 
England as well as the Eastern Townships and Beauce 
regions of Québec (Harrison and Chapin 1998; Mlade- 
noff and Sickley 1998; Carroll 2003*; Jolicoeur and 
Etcheverry, in preparation”). 

Wolves are highly mobile and frequently move over 
long distances (Van Camp and Glukie 1979; Fritts 
1983: Mech 1987). Some of the longest dispersal move- 
ments documented (460-555 km) occurred across a 
mixture of forest, farmland, and 4-lane highways in 
the upper Midwest (Mech et al. 1995; Wydeven et al. 
1995). A Wolf in Alberta crossed rivers 0.5 - 2.0 km 


' wide during summer (Van Camp and Glukie 1979), 
| and Wolves have crossed 24 km on a frozen lake (Mech 
1966). Wolves have increased in Minnesota (Fuller et 
) al. 1992), and since the mid-1970s have naturally re- 
colonized portions of northern Wisconsin and, more 
recently, Upper Michigan (Mech and Nowak 1981; 
Fuller et al. 1992). Wolves from Canada have recolo- 
nized Montana for at least the past decade (Boyd et al. 
1995). In a review of Wolf dispersal and recoloniza- 
| tion, Wydeven et al. (1998) reported that 31 to 63% 
of dispersing Wolves successfully settled and formed 
pairs in new territories. 

It is unclear whether Wolves have begun re-estab- 
_ lishing in the Eastern Townships, or whether this Wolf 
was simply a dispersing individual. While the Eastern 
Townships may be a sink habitat for Wolves, this Wolf 
was caught only 30-50 km from potential Wolf habi- 
tat in Maine (Carroll 2003*). Further investigation is 
necessary to confirm other possible Wolf observa- 
tions in these regions. Reporting of Wolf sightings or 
accidental Wolf captures south of the St Lawrence 
river should be encouraged. 


| Acknowledgments 

We are grateful to Laurent Cloutier, trapper and for- 
mer Eastern Townships Wildlife Protection Officer, 
for providing the specimen and details of the capture 
location. DNA analysis were funded by the Société 
de la Faune et des Parcs du Québec. Thanks to C. 
| Carroll, M. Festa-Bianchet, C. Reining and P. Struh- 


| sacker for comments on an earlier draft of this manu- 


| script. 


Documents Cited (marked * in text) 

Carrol, C. 2003. Carnivore restoration in the Northeastern 
U.S. and Southeastern Canada: a regional-scale analysis of 
habitat and population viability for wolf, lynx, and marten. 
Progress report 1: Wolf viability analysis. Prepared for 
The Wildlands Project. Orleans, California: Klamath Cen- 
ter for Conservation Research. 38 pages. 

Fortin, C., and J. Huot. 1995. Ecologie comparée du coyote, 
du lynx et du renard roux au pare national Forillon. Report 
prepared for Parks Canada. Département de Biologie, 
Universite Laval. Ste-Foy, Québec. 291 pages. 

_ Jolicoeur, H. 1998. Le loup du massif du lac Jacques-Cartier. 
Québec, Ministére de l’Environnement et de la Faune, 
Direction de la Faune et des Habitats, Direction de la 
conservation et du patrimoine écologique. 132 pages. 

Jolicoeur, H., and M. Hénault. 2002. Repartition géogra- 
phique du loup et du coyote au sud du 52° paralléle et 
estimation de la population de loups du Québec. Québec, 
Societe de la faune et des pares du Québec, Direction du 
développement de la faune et Direction de l’aménagement 
de la faune des Laurentides. 51 pages. 

| Jolicoeur, H., and P. Etcheverry (in preparation). Habitat 
potentiel pour le loup sur la rive sud du St-Laurent. Québec, 
Sociéte de la faune et des parcs du Québec, Direction du 
développement de la faune. 

Potvin, F. 1986. Ecologie du loup dans la reserve de Papi- 
neau-Labelle. Québec, Ministére du Loisir, de la Chasse 
et de la Péche, Direction générale de la Faune. 103 pages. 


NOTES 


609 


Villemure, M. 2003. Ecologie et conservation du Joup dans la 
region du parc national de la Mauricie. Mémoire de Maitrise. 
Département de Biologie, Universite de Sherbrooke. Sher- 
brooke, Québec, Canada. 89 pages. 


Literature Cited 

Boyd, D. K., P. C. Paquet, S. Donelon, R. R. Ream, D. H. 
Pletcher, and C. C. White. 1995. Transboundary move- 
ments of a recolonizing wolf population in the Rocky 
Mountains. Pages 141-146 in: Ecology and conservation 
of wolves in a changing world. Edited by L. N. Carbyn, 
S. H. Fritts and D. R. Seip. Canadian Circumpolar Insti- 
tute, Occasional Publication Number 35, 642 pages. 

Dumond, M., and M. A. Villard. 2000. Demography and 
body condition of coyotes (Canis latrans) in eastern New 
Brunswick. Canadian Journal of Zoology 78: 399-406. 

Fritts, S. H. 1983. Record dispersal by a wolf from Min- 
nesota. Journal of Mammalogy 64: 166-167. 

Fuller, T. K, W. E. Berg, G. L. Radde, M. S. Lenarz, and 
G. Blair Joselyn. 1992. A history and current estimate of 
wolf distribution and numbers in Minnesota. Wildlife 
Society Bulletin 20: 42-55. 

Harrison, D. J., and T. G. Chapin. 1998. Extent and con- 
nectivity of habitat for wolves in eastern North America. 
Wildlife Society Bulletin 26: 767-775. 

Lehman, N., A. Eisenhawer, K. Hansen, L. D. Mech, R. 
O. Peterson, P. J. P. Gogan, and R. K. Wayne. 1991. 
Introgression of coyote mitochondrial DNA into sympat- 
ric North American gray wolf populations. Evolution 45: 
104-119. ‘ 

Mech, L. D. 1966. The wolves of Isle Royale. U.S. National 
Park Service, Fauna Series 7. 

Mech, L. D. 1987. Age, season, distance, direction, and social 
aspects of wolf dispersal from a Minnesota pack. Pages 
55-74 in Mammalian dispersal patterns: The effects of 
social structure on population genetics. Edited by B. D. 
Chepko-Sade and Z. Tang Halpin. University of Chicago 
Press. Chicago. 342 pages. 

Mech, L. D., S. H. Fritts, and D. Wagner. 1995. Minnesota 

Wolf dispersal to Wisconsin and Michigan. American Mid- 

land Naturalist 133: 368-370. 

Mech, L. D., and R. M. Nowak. 1981. Return of the gray wolf 

to Wisconsin. American Midland Naturalist 105: 408-409. 

Messier, F. 1985. Solitary living and extraterritorial move- 

ments of wolves in relation to social status and prey abun- 

dance. Canadian Journal of Zoology 63: 239-245. 

Mladenoff, D. J., and T. A. Sickley. 1998. Assessing poten- 
tial gray wolf restoration in the northeastern United States 
—a spatial prediction of favorable habitat and potential 
population levels. Journal of Wildlife Management 62: 
1-10. 

Nowak, R. M. 1983. A perspective on the taxonomy of wolves 
in North America. Pages 10-19 in: Wolves in Canada and 
Alaska: their status, biology, and management. Edited by 
L. N. Carbyn. Canadian Wildlife Service. 145 pages. 

Nowak, R. M. 1995. Another look at wolf taxonomy. Pages 
375-397 in: Ecology and conservation of wolves in a chang- 
ing world. Edited by L. N. Carbyn, S. H. Fritts and D. R. 
Seip. Canadian Circumpolar Institute, Occasional Publi- 
cation Number 35, 642 pages. 

Nowak, R. M. 2002. The original status of wolves in eastern 
north america. Southeastern Naturalist 1: 95-130. 

Peterson, R. L. 1966. The mammals of eastern Canada. 
Oxford University Press. Toronto. 465 pages. 


610 


Potvin, F. 1987. Wolf movement and population dynamics 
in Papineau-Labelle reserve, Québec. Canadian Journal 
of Zoology 66: 1266-1273. 

Van Camp, J., and R. Glukie. 1979. A record long distance 
move by a wolf (Canis lupus). Journal of Mammalogy 60: 
236-237. 

Wayne, R K., N. Lehman, M. W. Allard, and R. L. Honey- 
cutt. 1992. Mitochondrial DNA variability of the gray wolf: 
Genetic consequences of population decline and habitat 
fragmentation. Conservation Biology 6: 559-569. 

Wilson, P. J., S. Grewal, I. D. Lawford, J. N. M. Heal, A. 
G. Granacki, D. Pennock, J. B. Theberge, M. T. The- 
berge, D. R. Voigt, W. Waddell, R. E. Chambers, P. C. 
Paquet, G. Goulet, D. Cluff, and B. N. White. 2000. 
DNA profiles of the eastern Canadian wolf and the red 
wolf provide evidence for a common evolutionary history 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


independent of the gray wolf. Canadian Journal of Zoology 
78: 2156-2166. 

Wydeven, A. P., T. K. Fuller, W. Weber, and K. Mac Donald. 
1998. The potential for wolf recovery in the Northeastern 
United States via dispersal from Southeastern Canada. 
Wildlife Society Bulletin 26: 776-784. 

Wydeven, A. P., R. N. Schultz, and R. P. Thiel. 1995. Moni- 
toring of a recovering gray wolf population in Wisconsin. 
Pages 147-156 in: Ecology and conservation of wolves 
in a changing world. Edited by L. N. Carbyn, S. H. Fritts, 
and D. R. Seip. Canadian Circumpolar Institute, Occa- 
sional Publication Number 35. 642 pages. 


Received 11 August 2003 
Accepted 24 December 2004 


Is Cost of Locomotion the Reason for Prolonged Nesting Forays of 
Snapping Turtles, Chelydra serpentina? 


SHANE R. DE SOLLA and KIM J. FERNIE 


Canadian Wildlife Service, Environment Canada, Canada Centre for Inland Waters, 867 Lakeshore Road, P.O. Box 5050, 
Burlington, Ontario L7R 4A6 Canada 


de Solla, Shane R., and Kim J. Fernie. 2004. Is cost of locomotion the reason for prolonged nesting forays of Snapping Turtles, 
Chelydra serpentina? Canadian Field-Naturalist. 118(4): 610-612. 


Prolonged nesting forays were observed in five gravid Snapping Turtles (Chelydra serpentina), in 1999 and 2001. For all 
observations, the females began exploratory nest excavations but failed to oviposit. Subsequently, all five females sought 
refuge either by burying themselves in substrate, or by seeking shade under vegetation, presumably to wait until the following 
day to resume nesting activities. By contrast, most observations of failed nesting resulted in the females abandoning the 
nesting site and returning immediately to water. Although prolonged nesting attempts in other turtle species (i.e. kinosternids) 
likely are associated with rainfall or predation risk, we speculate that these prolonged nesting attempts in Snapping Turtles 


reduced the cost of terrestrial travel. 


Key Words: Snapping Turtles, Chelydra serpentina, oviposition, nesting behaviour, cost of locomotion, Ontario 


Nesting behaviour of Snapping Turtles (Chelydra 
serpentina) has been described in detail elsewhere 
(see Ernst et al. 1994), and is generally characterized 
by behaviour that minimizes exposure on terrestrial 
habitats. Briefly, nesting behaviour starts with emer- 
gence from water, followed by nest site selection. 
Females typically dig with their front limbs, and often 
poke their noses into the substrate, presumably to deter- 
mine if the soil type and/or hydration are adequate. 
Subsequently, the rear limbs are used first to excavate 
the nest cavity, then position the eggs during oviposi- 
tion, and finally to bury the eggs. Subsequently, the 
female leaves the nesting area, and returns to the water. 
Although Snapping Turtles have been known to travel 
more than 0.5 km overland to nesting sites (Obbard 
and Brooks 1980), normally they nest at a site much 
closer to the water. Regardless if the female was suc- 
cessful in nesting or not, generally she returns to water 
immediately afterwards. Nesting normally occurs in 
the early morning or late evening, although females 
may nest in the afternoon following rain. By mini- 
mizing the distance or time traveled on land, female 
Snapping Turtles may avoid energetic costs, or the risks 


of dehydration or predation. Baudinette et al. (2000) 
found that terrestrial locomotion was 2.6 times more 
energetically costly than aquatic locomotion for the 
Murray Short-necked Turtle (Emydura macquarii). 
Snapping Turtles are particularly vulnerable to water 
loss compared to other turtles (Ernst 1968) due to their 
exposed skin. Although depredation of adults is rare, 
adult females are occasionally taken by large predators, 
such as bears or coyotes (Ernst et al. 1994). Here we 
document five instances of prolonged nesting forays 
in Snapping Turtles, which have not previously been 
documented, and discuss possible reasons for this 
behaviour. 


Study Site 

Observations were made from two locations in 
Ontario, both known for many years to be nesting sites 
for Snapping Turtles. The first nesting site, the north- 
western shoreline of Coote’s Paradise, Hamilton 
(43°16'N, 79°56'W), consists primarily of a communi- 
ty vegetable garden and a wood chip pile on Ontario 
Power Generation property. The nesting site is sepa- 
rated from the open water by approximately 150 m, 


first by a steep hill with thick shrub cover, and sec- 
-ondly by dense cattails. The second nesting site, at 


‘! Wheatley Provincial Park (42°5'N, 82°26'W), consists 


of a series of sand, dirt, and wood chip piles in a main- 
tenance area. Although normally open water is relative- 
ly close to the nesting site (~ 50 m), in 2001 when the 
observations were made, water levels were extremely 
low, and most water in the park had drained into Lake 
Erie, leaving bare mudflats. We observed many turtle 


| tracks in the exposed mud leaving the park into Lake 


Erie (de Solla, personal observation). Thus, at both 
sites, the nesting females did not have nearby access 
to water. 


| Nesting Observations 


De Solla et al. (2001) reported the deaths of two 
gravid Snapping Turtles that buried themselves in a 


| composting wood chip pile at Cootes Paradise, Hamil- 
ton, on 7 and 9 June 1999. Both gravid turtles remained 


in the wood chip pile until their deaths. Results of post- 
mortem examinations were consistent with death due 
to hyperthermia (de Solla et al. 2001); however, no 
reasons for their behaviour were suggested. The behav- 
iour of both females was consistent for females search- 


| ing for suitable nesting sites, except after failing to 
| successfully nest, both buried themselves in the wood 


chip pile. A third female Snapping Turtle was also 


| observed 8 June 1999 on the wood chip pile. Follow- 
| ing apparently unsuccessful attempts to nest, the female 
| was observed to be motionless for a few hours under 
| the shade of burdock (Arctium spp.) and other vege- 
| tation, which provided the only shade in the area. De 


Solla did not touch her until later in the morning, but 


| the turtle was found to be gravid. She remained in the 


shade after being disturbed. 

On 9 June 2001 at 1045, a female turtle was found 
on a storage area for sand, dirt, and wood chips at 
Wheatley Provincial Park. There were numerous tracks 
on both the wood chips and dirt piles, possibly by 
more than one female. There was abundant shade at 
this site. She was sitting in shade under vegetation on 
a small dirt pile, with no apparent movement, and no 
evidence of nesting. She was left alone until 1200, at 
which time the female was about 2 m from her previ- 
ous location, further in shade under vegetation. She 
did not move by 1300. At this time she was palpated, 
found to be still gravid, and a blood sample was taken. 
She was released at the same spot she was found, 
where she remained for at least one hour, but she was 
not subsequently observed. No temperature readings 
were taken; however, the weather was sunny and hot. 

On 12 June 2002, a female was seen in the vegetable 
garden at Coote’s Paradise at approximately 2130. 
Nearby were at least 10 digging attempts in freshly 
cultivated soil, and she was observed digging with 
her front limbs and poking her head into the soil. She 
was observed for approximately an hour, but did not 
show any signs of nesting. At approximately 2230, she 


NOTES 


611 


was observed to have buried herself, with only the top 
of her carapace showing. There was relatively little 
shade by vegetation available. At approximately 0745 
hrs, 13 June 2002, a female was seen walking very 
near where the female from the previous night was 
observed. This female was heavily covered with dirt, 
suggesting that she was the same female that had buried 
herself. No nesting attempts were made by 1000 hrs; 
subsequently we left and she was not observed again. 


Discussion 

The five observations of nesting attempts by Snap- 
ping Turtles had two common features; the delay was 
prior to oviposition, and the turtles sought terrestrial 
refuge by burying themselves in soil or wood chips, 
or by staying in shade. We have no observations of 
females remaining at the nesting ground after suc- 
cessful oviposition. We are unaware of any published 
reports of such behaviour in Snapping Turtles. 

Various kinosternid species prolong their nesting 
forays, typically by burying themselves before and/or 
after oviposition (Iverson 1990; Burke et al. 1994; Wil- 
son et al. 1999) adjacent to the nest. The duration of 
nest attendance in species other than Chelydra appears 
to be linked to the timing or duration of rainfall (Burke 
et al. 1994: Ernst et al. 1994). Intensive disturbance 
may also delay successful oviposition, and competitive 
nesting by Green Turtles (Chelonia mydas) at high 
densities may delay oviposition by a few days (Jessop 
et al. 1999). Typically, after a failed nesting attempt 
Snapping Turtles return to the wetland immediately 
following cessation of nesting behaviour (de Solla, 
persional observation). 

We speculate that the prolonged nesting attempts 
we observed in Snapping Turtles was to avoid the cost 
of making a second trip between the water and nest- 
ing site. All instances of prolonged nesting attempts 
that we observed occurred a considerable distance from 
water. Abandoning a nesting attempt would therefore 
result in a relatively long journey to the water, and 
back again to the nest site for a second oviposition 
attempt. To avoid this cost, once a female aborts a 
nesting attempt, she may burrow under substrate or 
hide under vegetation until the following day. Females 
nesting closer to wetlands are more likely to leave the 
nest site and come back at a later time. At the Wildlife 
Research Station (Algonquin Park), Snapping Turtle 
nesting has been monitored on a dam at the southern 
end of Lake Sasajewun for > 25 years, and all observed 
nesting attempts that failed were followed by the imme- 
diate withdrawal to the water (~ 5 m) by the female 
(R. J. Brooks, University of Guelph, personal com- 
munication). Similarly, at eight other nesting sites in 
Ontario that the authors visited, all of which were 
close to wetland habitat, females withdrew to water 
following a failed nesting attempt. We did not count 
the number of these observations, but de Solla has 
frequently observed this behaviour. 


612 


In two of the five observations of prolonged nesting 
attempts, the turtles died of hyperthermia after bury- 
ing themselves in composting wood chips (de Solla 
et al. (2001). We speculate that in most cases where 
females bury themselves, they use non-composting 
material, and thus are not at risk of hyperthermia. 


Acknowledgments 

We thank Wheatley and Algonquin Provincial Parks, 
and Ontario Power Generation for permission to enter 
their properties to look for turtle nests. Ron Brooks 
and anonymous reviewers gave valuable comments 
on earlier drafts. 


Literature Cited 

Baudinette R. V., A. M. Miller, and M. P. Sarre. 2000. 
Aquatic and terrestrial locomotory energetics in a toad and 
a turtle: a search for generalisations among ectotherms. 
Physiological and Biochemical Zoology 73: 672-682. 

Burke V. J., J. W. Gibbons, and J. L. Greene. 1994. Pro- 
longed nesting forays by common mud turtles (Kinosternon 
subrubrum). American Midland Naturalist 131: 190-195 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


de Solla S. R., D. Campbell, and C. A. Bishop. 2001. 
Hyperthermia induced mortality of gravid snapping tur- 
tles, Chelydra serpentina, and eggs in a wood chip pile. 
Canadian Field-Naturalist 115: 510-512. 

Ernst, C. H. 1968. Evaporative water-loss relationships of 
turtles. Journal of Herpetology. 2: 159-161. 

Ernst, C. H., J. E. Lovich, and R. W. Barbour. 1994. Turtles 
of the United States and Canada. Smithsonian Institution 
Press. Washington. 

Iverson, J. B. 1990. Nesting and parental care in the mud 
turtle, Kinosternon flavescens. Canadian Journal of Zoology 
68: 230-233. 

Jessop, T. S., C. J. Limpus, and J. M. Whittier. 1999. Plasma 
steroid interactions during high-density green turtle nest- 
ing and associated disturbance. General and Comparative 
Endocrinology 115: 90-100. 

Obbard M. E., and R. J. Brooks. 1980. Nesting migrations of 
the snapping turtle (Chelydra serpentina). Herpetologica 
36: 158-162. 

Wilson D. S., H. R. Mushinsky, and E. D. McCoy. 1999. 
Nesting behavior of the striped mud turtle, Kinosternon 
baurii (Testudines: Kinosternidae). Copeia. 1999: 958-968. 


Received 11 October 2003 
Accepted 23 November 2004 


Significant Vascular Plant Records from the Hamilton Area, Ontario 


CARL ROTHFELS 


Royal Botanical Gardens, 680 Plains Road W., Burlington, Ontario L7T 4H4 Canada 


Rothfels, Carl. 2004. Significant vascular plant records from the Hamilton area, Ontario. Canadian Field-Naturalist 118(4): 


612-615. 


Four additions to the known vascular flora of Ontario (Aesculus pavia, Ambrosia x helenae, Anthriscus caucalis, Verbena 
bonariensis) and ten other provincially significant records are discussed. Of the 14 taxa listed, two (Actaea x ludovici and 


Ambrosia X helenae) are native. 


Key Words: adventive species, flora, rare species, distribution, Hamilton, Halton, Ontario. 


Field and herbarium work centred around Royal 
Botanical Gardens on the border between Hamilton 
and Burlington, Ontario (43°29.00'N, 79°88.00'W), has 
resulted in the following 16 provincially significant 
vascular plant records in 2002. For locally significant 
records, see Rothfels (2003). All records are supported 
by specimens in the Royal Botanical Gardens herbar- 
ium (HAM). 

Taxa preceded by an asterisk (*) are not native to 
the Hamilton Region. Taxon names are followed by 
their subnational rank (Srank) as of 1998, where appli- 
cable (see Newmaster et al. 1998). An Srank of S1 
indicates a very rare native taxon, with generally five or 
fewer occurrences in Ontario; a rank of SE1 is similar, 
but refers to a non-native taxon; and a rank of SEH 
(historical) indicates a non-native species that has 
typically not been recorded in Ontario in the past 20 
years (Newmaster et al. 1998). The names used follow 
Kartesz and Meacham (1999*). 


Actaea x ludovici B.Boivin. Hybrid Baneberry S1 
RANUNCULACEAE 

One clump of this hybrid was collected from the 
Devil’s Punchbowl Environmentally Significant Area, 
in Hamilton (43°11.80'N, 79°38.50'W). Both parental 
species (Actaea rubra (Aiton) Willd. and Actaea 
pachypoda Ell.) were in close proximity. Despite the 
overlapping ranges of the parental taxa, the hybrid is 
rare (Voss 1985), perhaps due to differing phenology 
(Pringle, personal communication). In Canada, it is 
recorded from Ontario and Quebec (Kartesz and 
Meacham 1999*). (C. J. Rothfels 305). 


* Aesculus pavia L. Red Buckeye HIPPOCASTANACEAE 

This species is an addition to the flora of Ontario 
(Newmaster et al. 1998). Over ten young trees were 
found naturalized in the Red Hill Creek escarpment 
valley, City of Hamilton, by Megan Ogilvie and Albert 
Garofalo. These plants are likely spreading from trees 


2004 


| planted in 1927 (Bruce Duncan, personal communica- 
) tion). A species of the southern United States, it 
| reaches its northern limit in Kentucky, West Virginia, 
and southern Illinois (Kartesz and Meacham 1999*),. 
| It differs from A. glabra Willd. and A. hippocastanum 
| L., the other two species reported from Ontario, by its 
/ smooth fruit (Edmondson 1997), among other features. 
| It is, due to its reddish flowers, perhaps most likely to 
| be confused with the frequently-planted Aesculus 
_ xcarnea Hayne, which differs in its size (A. pavia is a 
| small tree or large shrub, A. xcarnea is a large tree), 
_ leaf shape (A. pavia has lanceolate leaves unlike the 
| obovate leaves of A. xcarnea) and petal shape (A. 
_ pavia has dissimilar petals with two broad lateral 
_ petals and two long-stalked spoon-shaped petals; all 
_ petals of A. xcarnea are large and shaggy) (Krussman 
_ 1984). (M. Ogilvie, A .Garofalo et al. s.n.; HAM 16221). 


_ * Aethusa cynapium L. Fool’s-parsley SE1 APIACEAE 

This species was found new for both the City of 
Hamilton (Goodban 1995) and Lambton County (Tiedje 
and Tiedje 2002) in 2002. In Hamilton, it is fairly 
common along wooded paths along the base of the 
Niagara Escarpment in the Devil’s Punchbowl Envi- 
ronmentally Significant Area (43°12.20'N, 79°40.70'W). 
_ The Hamilton specimen was determined only to the 
_ specific level. The Lambton County specimen was 
_ determined as A. cynapium ssp. cynapioides (M. Bieb.) 
_ Nyman, the larger woodland subspecies (using Tutin 
1968) and it was common on the Ausable River flood- 
plain in the Rock Glen Conservation Area (43°05.10'N, 
81°06.80'W). Aethusa cynapium has been found in the 
Great Lakes States, and in Quebec, New Brunswick, 
and Nova Scotia (Kartesz and Meacham 1999*). The 
early Canadian reports describe this species as a garden 
weed “introduced with garden seeds from Europe” 
(Macoun 1883), and list it from Hastings and North- 
umberland Counties in Ontario, and as “occasional” 
in New Brunswick (Macoun 1883). (J. Rothfels & J. 
Shearer 243; C. J. Rothfels 274). 


* Amaranthus blitum L. Purplish Amaranth SEH 
AMARANTHACEAE 

This species is new for Halton Region (Varga et al. 
2000). A purple-leaved amaranth, it is scattered irreg- 
ularly in the lawns at Royal Botanical Gardens Centre 
(43°17.40'N, 79°59.70'W) where it appears to be dis- 
persed by Canada Geese. Scoggan (1978) reports this 
species as introduced in waste ground in Elgin County, 
Huron County, and Waterloo Regional Municipality 
in Ontario, and from Masson, Montreal and Quebec 
City in Quebec. (C. J. Rothfels & D. Gugler 303). 


Ambrosia x helenae Rouleau Hybrid Ragweed 
ASTERACEAE 

This taxon is an addition to the flora of Ontario 
(Newmaster et al. 1998). It is a hybrid between Com- 
mon Ragweed (Ambrosia artemisiifolia L.) and Giant 
Ragweed (Ambrosia trifida L.). Although the ranges 


NOTES 


613 


of these two species overlap frequently, the hybrid is 
very rare (Wagner 1958). It was collected along the 
north shore trails of the Cootes Paradise Nature Sanc- 
tuary in Hamilton (43°16.90'N, 79°54.10'W). The 
type locality is in Quebec (Wagner 1958), and it has 
also been found in Michigan, New York State, and 
Ohio (Kartesz and Meacham 1999*). (D. Gugler s.n.: 
HAM 15959). 


* Anthriscus caucalis M. Bieb. Bur-chervil APIACEAE 

This species is an addition to the flora of Ontario 
(Newmaster et al. 1998). It was abundant on disturbed 
ground near the Royal Botanical Gardens’ Laking 
Garden (43°17.50'N, 79°53.30'W). Shortly after the 
discovery, the site was mowed by the owner. This 
species is known in Canada only from Nova Scotia, but 
is scattered widely across the United States (Kartesz 
and Meacham 1999*), and is easily distinguished from 
its more common congener (A. sylvestris (L.) Hoffm.) 
by the hooked hairs on the fruit (e.g., Gleason and 
Cronquist 1991). (C. J. Rothfels & J. L. Reader 115). 


* Anthyllis vulneraria L. Lady’s-fingers SE1 
FABACEAE 

This species is new for the City of Hamilton (Good- 
ban 1995). It was fairly common, but local, in disturbed, 
weedy, poorly drained ground (a pipeline right of way) 
in the Beverly Swamp Environmentally Significant 
Area (43°21.10'N, 80°06.80'W). It has been reported 
from British Columbia, Ontario, Quebec, Newfound- 
land, and New Brunswick (Kartesz and Meacham 
1999*). Scoggan (1978) notes that it is “locally intro- 
duced into clover fields and waste places,” a description 
that concords closely with this report, and lists records 
from Oxford, Waterloo and Wellington Counties in 
Ontario, from “slaty banks of the Restigouche River 
near Matapedia” in Quebec, and from Newcastle in 
New Brunswick. (C. J. Rothfels 143). 


* Cardamine impatiens L. Narrow-leaved Bittercress 
SE1 BRASSICACEAE 

This species is new for the City of Hamilton (Good- 
ban 1995). Four plants were found along the north 
shore of the Cootes Paradise Nature Sanctuary, near a 
small boat-storage facility, along a partially disturbed 
forest edge (43°16.90'N, 79°54.00'W). This is an up- 
right Cardamine, with distinctly auriculate pinnately- 
divided leaves (e.g. Voss 1985). Mulligan (2002) only 
saw a single Canadian specimen, collected by J. M. 
Weber in 1980 for Port Credit, Ontario, and notes that 
this species is “sporadic and uncommon” in the United 
States. This species was not listed by Scoggan (1978). 
(C. J. Rothfels 88). 


* Coronopus didymus (L.) J.E.Sm. Lesser Wart-cress 
SEI BRASSICACEAE 

This strange little mustard is new for the City of 
Hamilton (Goodban 1995). It is a common weed in 
some of the Royal Botanical Gardens’ Rock Garden 


614 


flower beds (43°17.20'N, 79°53.60'W). This is the 
second confirmed record for Ontario (Oldham personal 
communication). It first established a foothold in the 
east — Scoggan (1978) lists five counties in Nova 
Scotia and two in New Brunswick containing this 
species — but it also occurs across the country (a single 
record each in British Columbia, Alberta, Quebec, and 
Newfoundland (Scoggan 1978). The first Canadian 
records are from Gaspé Basin, Quebec, and North 
Sidney, Nova Scotia, in 1862 and 1883, respectively 
(Mulligan 2002). Coronopus didymus can be differen- 
tiated from Coronopus squamatus (Forssk.) Aschers. 
(which is also weedy) by its wrinkled cordate fruits that 
are notched at the summit. The fruits of C. sguamatus 
are conspicuously apiculate at the summit (e.g. 
Holmgren 1998). (C. J. Rothfels 71). 


* Euonymus fortunei (Turcz.) Hand.-Mazz. Wall- 
creeper SE] CELASTRACEAE 

This species has been known from Royal Botani- 
cal Gardens’ sanctuary lands for several years (Smith 
2003), but has not been collected until this year. It is 
widespread but local in the Cootes Paradise Nature 
Sanctuary, where it occasionally forms small dense 
patches on the oak-hickory slopes (43°1620'N, 
79°55.10'W). It seems to be increasing in southern 
Ontario, and may become a problem in some natural 
areas (Oldham, personal communication 2003). (C. 
J. Rothfels 235). 


* Macleaya cordata (Willd.) R. Br. Plume-poppy 
SEH PAPAVERACEAE 

This species is new for the City of Hamilton (Good- 
ban 1995). This large plant is occasionally planted as 
an ornamental. Its occurrence on the east shore of the 
Cootes Paradise Nature Sanctuary (43°16.80'N, 
79°53.50'W) is mysterious; it could be a persistent 
population from some unknown source or it could be 
“spontaneous”. Voss (1985) describes it as “seldom 
escaping” in Michigan, and Scoggan (1978) lists it as a 
“garden escape” that is “scarcely established” in Nor- 
folk County in Ontario and Missisquoi County in 
Quebec. (C. J. Rothfels 318). 


* Myrrhis odorata (L.) Scop. Scented Myrrhis SE1 
APIACEAE 

This large umbellifer is new for Halton Region 
(Varga et al. 2002). It is escaping from the Royal Bot- 
anical Gardens’ Scented Garden into the neighboring 
ravines (43°17.50'N, 79°52.60'W). Currently, its spread 
is limited, and will be tracked in future years. It is 
reported from two other Ontario locations: St. Thomas, 
Elgin County, and Manitoulin Island, Manitoulin 
District (Pringle 1994); as well as from British Colum- 
bia, Nova Scotia, Michigan, Pennsylvania, and Oregon 
(Kartesz and Meacham 1999*). The smooth fruit and 
large size help differentiate this species from the native 
Osmorhiza species (Pringle 1994). (C. J. Rothfels 
190). 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


* Sorghum bicolor (L.) Moench ssp. bicolor. Sorghum 
SE] POACEAE 

This species is new for the City of Hamilton (Good- 
ban 1995). It was common with other weeds on freshly 
disturbed ground at the Royal Botanical Gardens’ 
outdoor compost facility (43°17.10'N, 79°53.70'W). 
It has been found in Quebec and most U.S. states, 
and is a noxious weed in three states (Kartesz and 
Meacham 1999). (C. J. Rothfels 317). 


* Verbena bonariensis L. Purpletop Vervain VERBE- 
NACEAE 

This species is an addition to the flora of Ontario 
(Newmaster et al. 1998). One plant was found on the 
Osprey Marsh Christmas tree carp barrier (a row of 
old Christmas trees erected in the mud to prevent carp 
from travelling from Grindstone Creek into Osprey 
Marsh), in the Hendrie Valley Nature Sanctuary, Halton 
Region (43°17.40'N, 79°52.90'W). It might have ar- 
rived in the treads of the machinery used to place the 
trees in the winter. This species is not in the Royal 
Botanical Gardens’ cultivated plants database, and 
thus has theoretically never been planted on RBG 
property. One individual of this species was seen at 
the Childrens’ Garden in Westdale (not operated by 
RBG) in 2002, approximately 3.5 km from the Osprey 
Marsh population. This species is introduced to the 
southern States, and has also been reported from Ore- 
gon, New York State, New Jersey, and New Hampshire 
(Kartesz and Meacham 1999*). It has not been re- 
ported from Canada. (C. J. Rothfels & I. Vaithilingam 
435). A previously-overlooked specimen was also 
uncovered at HAM. It is an undated specimen from a 
“river bank” at “Lake Erie,’ and was identified by J. 
S. Pringle in 1964 (K. Stanley s.n.). In light of the 
2002 discovery, it seems reasonable that this record 
could also have been “spontaneous”. 


Discussion 

The eight taxa found new for Hamilton add to the 
current published list of 1304 species (Goodban 1995), 
and the four new taxa for Halton bring that region’s 
tally to 1305 (Varga et al 2000). Additionally, four of 
the taxa are new for Ontario. Twelve of the fourteen 
records discussed are non-native occurrences, several 
of which, especially Aethusa cynapium, Euonymus 
fortunei, and Sorghum bicolor, should be watched for 
invasive tendencies. 


Acknowledgments 

Thanks to Jim Pringle for assistance in the deter- 
mination of Aethusa cynapium, Amaranthus blitum, 
Anthriscus caucalis, Cardamine impatiens, Coronopus 
didymus, and Macleaya cordata; thanks to Jim Pringle 
and Michael J. Oldham for helpful earlier reviews of 
this manuscript, and to Megan Ogilvie, Albert Garofalo 
and Dean Gugler for submitting their records. Con- 
tribution from Royal Botanical Gardens Number 1 14. 


2004 NOTES 615 
Documents Cited (marked * in text) Pringle, J. S. 1994. Notes on some adventive and naturalized 
Kartesz, J. T., and C. A. Meacham. 1999. Synthesis of the species in the flora of Ontario. Field Botanists of Ontario 
North American Flora, Version 1.0. Chapel Hill: North Newsletter. 7(4): 10-14. 
Carolina Botanical Garden. Compact disc. Rothfels, C. J. 2003. Significant 2002 Hamilton Study Area 


Literature Cited: 

Edmondson, J. R. 1997. Aesculus. Pages 155-157 in The 

/ European garden flora Volume V. Edited by J. Cullen, J. 
C. M. Alexander, C. D. Brickell, J. R. Edmondson, P. S. 
Green, V. H. Heywood, P. —M. Jorgensen, S. L. Jury, S. 
G. Knees, V. A. Matthews, H. S. Maxwell, D. M. Miller, 
E. C. Nelson, N. K. B. Robson, S. M. Walters, and P. F. 
Yeo. Cambridge University Press, United Kingdom. 

Gleason, H. A., and A. Cronquist. 1991. Manual of vascular 
plants of Northeastern United States and adjacent Canada. 
New York Botanical Garden. Bronx, New York. 910 pages. 

_ Goodban, A. G. 1995. The vascular plant flora of the Regional 
Municipality of Hamilton-Wentworth, Ontario. First edition: 
September 1995. Hamilton Region Conservation Authority. 
86 pages. 

Holmgren, N. H. 1998. Illustrated Companion to Gleason and 
Cronquist’s Manual. New York Botanical Garden. Bronx, 
New York. 937 pages. 

Krussmann, G. 1984. Manual of cultivated broad-leaved 
trees & shrubs. B. T. Batsford Ltd., London. 448 pages. 
Macoun, J. 1883. Catalogue of Canadian plants: Part 1. 

Geological Survey of Canada. Ottawa. 

Mulligan, G. A. 2002. Weedy introduced mustards (Bras- 
sicaceae) of Canada. Canadian Field-Naturalist. 116: 623- 
631. 

Newmaster, S. G., A. Lehela, P. W. C. Uhlig, S. McMurray, 
and M. J. Oldham. 1998. Ontario Plant List. Ontario 
Ministry of Natural Resources, Ontario Forest Research 
Institute, Sault Ste. Marie, Ontario, Forest Research 
Information Paper Number 123, 550 pages + appendices. 


Plant Records from the Royal Botanical Gardens Herbar- 
ium (HAM). Wood Duck 56: 155-161. 

Scoggan, H. J. 1978. The flora of Canada. National Museums 
of Canada. Ottawa. Four Parts; 1711 pages. 

Smith, T. W. 2003. Checklist of the Vascular Flora of Royal 
Botanical Gardens. Contribution of Royal Botanical Gar- 
dens Number 113. Royal Botanical Gardens, Burlington, 
Ontario. 126 pages. 

Tiedje, J., and D. Tiedje. 2002. Vascular plants of Lambton 
County, Ontario. 11' Edition: November 2002. Published 
by the authors. 66 pages. 

Tutin, T. G. 1968. Aethusa. Pages 339-340 in Flora Europaea 
Volume 2. Edited by T. G. Tutin, V. H. Heywood, N. A. 
Burges, D. M. Moore, D. H. Valentine, S. M. Walters, 
and D. A. Webb. Cambridge University Press. 455 pages. 

Varga, S., D. Leadbeater, J. Webber, J. Kaiser, B. Crins, J. 
Kamstra, D. Banville, E. Ashley, G. Miller, C. Kingsley, 
C. Jacobsen, K. Mewa, L. Tebby, E. Mosley, and E. 
Zaje. 2000. Distribution and status of the vascular plants 
of the greater Toronto Area. Ontario Ministry of Natural 
Resources, Aurora District. August 2000. 102 pages. 

Voss, E. G. 1985. Michigan Flora: Part II Dicots (Saururaceae 
to Cornaceae). Cranbrook Institute of Science Bulletin 
59. University of Michigan Herbarium. 724 pages. 

Wagner, W. H., Jr. 1958. The hybrid ragweed, Ambrosia 
artemisiifolia x trifida. Rhodora 60: 309-316. 


Received 10 November 2003 
Accepted 21 December 2004 


Do Rhinoceros Auklet, Cerorhinca monocerata, Fledglings Fly to the 


Sea from Their Natal Burrows? 


JAMES L. HAYWARD and JERE K. CLAYBURN 


Biology Department, Andrews University, Berrien Springs, Michigan 49104-0410 USA 


Hayward, James L., and Jere K. Clayburn. 2004. Do Rhinoceros Auklet, Cerorhinca moncerata, fledglings fly to the sea 
from their natal burrows? Canadian Field-Naturalist 118(4): 615-617. 


The mode of departure of Rhinoceros Auklet fledglings from their nest burrows has remained uncertain. Both walk-down 
and fly-down hypotheses have been proposed. Here we use the unique geography of Protection Island, Washington, to eval- 
uate the fly-down hypothesis. Some fledglings raised on Protection Island do appear to walk to the water, but our results 
suggest that many of the island’s fledgling Rhinoceros Auklets fly to the sea. 


Key Words: Rhinoceros Auklet, Cerorhinca monocerata, fledgling departure, Washington. 


Rhinoceros Auklets (Cerorhinca monocerata) are 
the most abundant puffins in the waters of the Pacific 
Northwest. Unlike Tufted and Horned puffins (Frater- 
cula cirrhata and F: corniculata, respectively), which 
also breed along western North America, members of 
most populations of Rhinoceros Auklets leave their 
nesting colonies before dawn to feed and return after 
sunset (Wilson and Manuwal 1986; but see Thoresen 
1980). Consequently, some basic information on the 


behavior of these birds remains obscure, despite grow- 
ing knowledge of their breeding biology (Richardson 
1961; Scott et al. 1974; Leschner 1976; Summers and 
Drent 1979; Vermeer 1978, 1979, 1980; Thoresen 1980, 
1983; Wilson and Manuwal 1986; Wilson 1993; Gaston 
and Duchesne 1996). 

One elusive piece of information involves the mode 
of departure from nest sites by newly-fledged Rhinoc- 
eros Auklets. Young Atlantic Puffins (F. arctica) walk, 


616 


flutter, or fly to the water from their burrows (Lockley 
1934: Richardson 1961; Gaston and Jones 1998), where- 
as fledgling Cassin’s Auklets (Ptychoramphus aleuti- 
cus) fly to the sea after making short “practice flights” 
(Manuwal 1974). Leschner (1976) found Rhinoceros 
Auklet fledglings that were too small to fly “crouched 
under rocks or crevices in the morning” on Destruction 
Island, Washington; this suggested to her “that fledg- 
lings walk or flutter to the water.” Similarly, Wilson 
(1977) observed fledgling Rhinoceros Auklets on Pro- 
tection Island, Washington, “walking down the grass 
covered slopes” to the sea. But Richardson, (1961) who 
also worked on Protection Island, wrote that “flight 
[capability] ... appears to develop precociously,’ and 
that like the “half-grown young” of murres, fledgling 
Rhinoceros Auklets “fly down to the water.” 

On Protection Island, hundreds of Rhinoceros Auk- 
lets nest along a sandy cliff that rises above Violet Point, 
a gravel spit that contains a large Glaucous-winged 
Gull (Larus glaucescens) colony. Each summer, scores 
of auklet fledglings are found dead within a relatively 
defined area of the gull colony, across an artificial 
channel and marina from the cliff. The distribution of 
dead fledglings and the unique semi-natural geography 
of Protection Island allowed us to evaluate Richard- 
son’s fly-down hypothesis. 


Methods 

Protection Island, Jefferson County, Washington 
(48°08'N, 122°55'W), consists of a 35—76-m high pla- 
teau surrounded by steep, grass-covered, sandy cliffs 
ideally suited for Rhinoceros Auklet burrowing. The 
largest Rhinoceros Auklet colony in Washington is 
located here, with approximately 27 549 burrows (Wil- 
son and Manuwal 1986). Violet Point, a gravel spit that 
extends 800 m east from the upper island, contained 
around 5 100 of Glaucous-winged Gull nests during 
this study (J. G. Galusha, personal communication). 

In July and August 1992, 1993, and 1997 the location 
of each auklet fledgling found dead on Violet Point was 
plotted by measuring its north or south distance from 
a transect that measured its eastward distance from the 
base of the island’s east-facing cliff. To estimate age at 
death, culmen and tarsal lengths, if still present, were 
measured in 1992 and 1993. The height of the highest 
(60 m) nesting burrows on the cliff in relation to the 
distribution of dead fledglings on Violet Point was used 
to approximate a range of presumed fledgling flight 
path angles trigonometrically. 


Results 

One hundred and twenty-one dead fledglings (43 in 
1992, 11 in 1993, and 67 in 1997) were counted between 
106 m east of the cliff base, at the western margin of the 
gull colony, and 398 m east of the cliff base, toward 
the center of the gull colony (Figure 1). The fledglings 
apparently died as a result of hitting the spit or being 
attacked by resident gulls. No dead fledglings were 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


A. Aerial view 


Strait of 
Juan de Fuca 


Protection 
Island 


60-m cliff 
with auklet 
burrows 


Violet Point 


(Exeter Of quill COL Cy) 
_—_—_—_——_——_——[$ (478 m - longest estimated fledgling flight) 
See! (111 m - shortest estimated fledgling flight) 
bt (292 m - range Of dead fledglings) 


B. Side view 


Ficure |. A. Aerial view of Violet Point, Protection Island, 
Washington. Dots indicate locations of dead fledgling 
Rhinoceros Auklets found during July and August, 1992, 
1993, and 1997. Lines below indicate approximate extent 
of the spit occupied by the gull colony, extent of fledgling 
auklet corpses, and estimated extents of fledgling auklet 
flight distances. B. Side view of Violet Point showing the 
estimated range of Rhinoceros Auklet fledgling flight 
paths, sloping from 7°—18° from the highest (60 m) natal 
burrows above the gull colony; fledgling flights from 
lower burrows would exhibit progressively shallower 
angles. 


found beyond 398 m. The pooled mean (+ SD here 
and below) distance of fledgling corpses from the base 
of the nest cliff was 303 + 58.0 m (n = 54, range = 
106-398 m). Mean culmen length was 24.4 + 3.0 mm 
(n = 35), and mean tarsus length was 37.9 + 1.6 mm 
(n = 48); Rhinoceros Auklets of this size are about 
45 days old (Wilson and Manuwal 1986). During late 
July and early August, we occasionally found live 
Rhinoceros Auklet fledglings hiding in grass between 
the nesting cliff and the channel/marina, and we reg- 
ularly observed fledglings swimming in the water of 
the marina/channel. 


Discussion 

Large numbers of dead fledglings found in gull ter- 
ritories east of the marina/channel water and across 
from the nesting cliff support the hypothesis that these 
fledglings flew down and over the water from their 
natal burrows. Alternate means of reaching the east 
side of the marina/channel were highly unlikely: (1) 
walk/flutter down slope, enter the marina/channel 
water from the west shore, leave the water on the east 
shore, enter the gull colony, be killed; (2) walk/flutter 


2004 


down slope, enter the colony north of the marina/ 
channel, walk/flutter through a gauntlet of gull terri- 
tories to the east side of the water, be killed; (3) walk/ 
flutter down slope, enter the colony north of the 
marina/channel, be killed, then be transported to dis- 
tant points east of the water by gulls. With respect to 
(3), it should be noted that fledging Rhinoceros Auk- 
lets exhibit an average mass of 360 g (Wilson and 
Manuwal 1986), 34% of the average mass of 1051 g 
(Vermeer 1963) for adult Glaucous-winged Gulls in 
this region; thus it appears unlikely that auklet corpses 
could be moved very far from the place where they 
were killed. 

Our data, with those of Richardson (1961), Lesch- 
ner (1976), Wilson (1977), and Wilson and Manuwal 
(1986), support the following scenario for the depar- 
ture of fledgling Rhinoceros Auklets from their nest 
burrows on Protection Island: In late July and early 
August, some fledgling auklets walk and flutter, but 
also many fly directly away from their natal burrows. 
Fledglings cannot sustain prolonged flight, however, 
and from the highest nests lose altitude at a rate of 
13-32 m/100-m distance, with a slope of 7°-18° 
(Figure 1B); fledgling flights from lower burrows 
would exhibit progressively shallower angles. Fledg- 
lings from preferred nest sites located directly above 
the ocean (Richardson 1961) reach the water success- 
fully. Fledglings that fly from nest sites along the cliff 
above Violet Point hit the gull colony where they are 
killed, unless they veer to the ocean or are fortunate 
enough to land in the marina/channel. Whether this 
scenario applies to fledgling departures at other Rhi- 
noceros Auklet colonies remains unknown. 


Acknowledgments 

We thank Robert Edens, Jean Takekawa and Kevin 
Ryan for permission to work on Protection Island 
National Wildlife Refuge, Ulrich Wilson for valuable 
discussions about Rhinoceros Auklets, Shandelle Hen- 
son for comments on the manuscript, and Walla Walla 
College Marine Station for logistical support during 
this study. Funding was provided by faculty grants to 
JLH from Andrews University. 


Literature Cited 
Gaston, A. J., and S. B. C. Dechesne. 1996. Rhincoceros 
Auklet: Cerorhinca monocerata. The Birds of North Amer- 


NOTES 


617 


ica (212). American Ornithologists’ Union, Washington, 
D. C., and Academy of Natural Sciences, Philadelphia, 
Pennsylvania. 20 pages. 

Gaston, A. J., and I. L. Jones. 1998. The Auks: Alcidae. 
Oxford University Press, Oxford. 349 pages. 

Leschner, L. L. 1976. The breeding biology of the Rhinoceros 
Auklet on Destruction Island. M.S. thesis, University of 
Washington, Seattle. 77 pages. 

Lockley, R. M. 1934. On the breeding habits of the puffin: 
With special reference to its incubation and fledgling peri- 
ods. British Birds 27: 214-223. 

Manuwal, D. A. 1974. The natural history of Cassin’s Auk- 
let (Ptychoramphus aleuticus). Condor 76: 421-431. 

Richardson, F. 1961. Breeding biology of the Rhinoceros 
Auklet on Protection Island, Washington. Condor 63: 456— 
473. 

Scott, J. M., W. Hoffman, D. Ainley, and C. F. Zeillemaker. 
1974. Range expansion and activity patterns in Rhinoceros 
Auklets. Western Birds 5: 13-20. 

Summers, K. R., and R. H. Drent. 1979. Breeding biology 
and twinning experiments of Rhinoceros Auklets on Cle- 
land Island, British Columbia. Murrelet 60: 16-22. 

Thoresen, A. C. 1980. Diurnal land visitations by Rhinoceros 
Auklets. Western Birds 11: 154. 

Thoresen, A. C. 1983. Diurnal activity and social displays of 
Rhinoceros Auklets on Teuri Island, Japan. Condor 85: 373— 
SU 

Vermeer, K. 1963. The breeding ecology of the Glaucous- 
winged Gull (Larus glaucescens) on Mandarte Island, 
B. C. Occasional Papers of the British Columbia Provin- 
cial Museum Number 13. 104 pages. 

Vermeer, K. 1978. Extensive reproductive failure of Rhi- 
noceros Auklets and Tufted Puffins. Ibis 120: 112. 

Vermeer, K. 1979. Nesting requirements, food and breeding 
distribution of Rhinoceros Auklets, Cerorhinca monocer- 
ata, and Tufted Puffins, Lunda cirrhata. Ardea 67: 101— 
110. 

Vermeer, K. 1980. The importance of timing and type of prey 
to reproductive success of Rhinoceros Auklets Cerorhinca 
monocerata. Ibis 122: 342-350. 

Wilson, U. W. 1977. A study of the biology of the Rhinoceros 
Auklet on Protection Island, Washington. M.S. thesis, Uni- 
versity of Washington, Seattle. 98 pages. 

Wilson, U. W. 1993. Rhinoceros Auklet burrow use, breeding 
success, and chick growth: Gull-free vs. gull-occupied habi- 
tat. Journal of Field Ornithology 64: 256-261. 

Wilson, U. W., and D. A. Manuwal. 1986. Breeding biology 
of the Rhinoceros Auklet in Washington. Condor 88: 143— 
156. 


Received 6 February 2004 
Accepted 25 October 2004 


A Tribute to Thomas Henry Manning 1911-1998 


BRENDA CARTER 


R.R. #4. Merrickville, Ontario KOG 1NO Canada 


Carter, Brenda. 2004. A tribute to Thomas Henry Manning 1911-1998. Canadian Field-Naturalist 118(4): 618-625. 


I first met Tom Manning on his Merrickville farm 
in the spring of 1965, during lambing time. Andrew 
and Betty Macpherson who had both travelled with 
him in the Arctic, brought me out to see Arctic Foxes. 
These captive animals were part of a research program 
on the influence of light on moult timing. On the way 
to. Merrickville, I listened to their stories about Tom’s 
legendary explorations and scientific expeditions. I was 
therefore prepared for the prospect of meeting an Arc- 
tic hero. Instead, I found a quiet, stocky man in his 
fifties, dressed in faded and ragged overalls tending 
his sheep. The stub of a hand-rolled cigarette hung from 
his lower lip — perpetually unlit. He did not seem pleased 
at extra company and bluntly asked why I had been 
brought out. Later, over glasses of his lethal home- 
made cider, I realized that here was a man who was 
much more than he seemed. By the end of that day we 
had become firm friends — a friendship that was to last 
until his death in November of 1998. 

Few knew Tom Manning well; and few indeed were 
given a chance to comprehend, as a whole, the com- 
plexity and sheer bulk of his phenomenal life’s work. 
Except for a very few close friends, Tom was deeply 
private, and even withdrawn. He was a relentless and 
dogged scientist who spared no effort to find either 
the original source of any information, or to redo any 
questionable measurements — even if they were his 
own. Applying his logical and inquiring mind to all 
kinds of life-forms, natural phenomena and expedi- 
tion challenges, Tom also had the skill, intuition and 
knowledge to assess and react appropriately in critical 
situations. He did so with a rapidity I found astonishing. 
His mental brilliance, awesome physical endurance, 
good looks and charm, combined with great modesty 
and fondness for red sportscars, contributed to a re- 
markable personality. 

On the 22 of December, 1911, Thomas Henry Man- 
ning was born into a gentleman’s life. Even as an infant, 
he defied prediction and showed his amazing capacity 
for endurance. Except for a surgeon who “did not want 
to commit murder on Christmas Day” and reluctantly 
operated, Tom might soon have died, a victim of pyloric 
stenosis. Tom was the only child of Dorothy (nee Ran- 
dall) whose brother was the last gentleman jockey in 
England, and Thomas Edward Manning, (TEM) who 
managed the family brewery and was cricket Captain 


for Northamptonshire. Their privileged son was neither 
destined nor inclined to follow in his father’s foot- 
steps. 

Tom once told me that his first childhood memory 
was of being put out on a blanket in the sun before he 
could walk. He remembered his excitement and free- 
dom at being outside. Tom never lost his passion for 
open air which propelled him into a life of exploration 
and adventure. His scientific career began and ended 
with eggs. As a boy, he had a passion for collecting eggs, 
and this hobby directed him into a career as a biologist, 
rather than a brewery owner as his father expected. A 
friendly gardener sparked his interest in nests, and Tom 
undoubtedly learned to keep notes and data for his tro- 
phies. This mixture of intense interest and attention 
to detail stood him in good stead as he completed, at 
the age of seventy, his last pioneering scientific work on 
weight lost by eggs during incubation for the Canadian 
Field-Naturalist. 

Tom was educated at Harrow, and like his father, 
attended shooting parties and rode with the famous 
Pytchley Hunt. There were, however, some interesting 
dissimilarities. He cleaned the skulls of the mammals 
and birds shot, drying them in his mother’s linen cup- 
board. Eventually, he escaped from the hunt meets be- 
cause he disliked the formalities and confining clothes. 

While at Harrow, Tom took a cycling trip with a 
school friend and discovered an unsuspected physical 
trait. As the pair wound their way through country 
lanes toward the southern coast of England, he found 
himself farther and farther ahead. With characteristic 
independence and detachment from social niceties, 
he left his friend far behind and continued on, solely 
interested in how far he could travel in one day. At the 
end of that journey, he was surprised by his endurance. 
Although he would never admit it, this ability was 
more than unusual. 

After his death, I discovered a buried manuscript, 
transcribed from a 1931 diary. It gives an account of 
his first expedition to Iceland and the Faroe Islands 
with Harrow school friend, “Twitch” Mitchell. Tom’s 
descriptions of unidentified gulls and seabirds revealed 
how scrupulous his note-taking was at the young age 
of twenty. Equally remarkable, for a twenty-year old, 
was the care he took in logistical planning; from 
deciding on exact food supplies and arranging trans- 


618 


CARTER: THOMAS HENRY MANNING 


Thomas Henry Manning with Massey Medal presented by the Royal Canadian Geographical Society in 1977 with portrait 
of former Government-General Vincent Massey in background. 


portation, to sorting out necessary visas. This thor- pithy comments about the unfortunate “Twitch”, dis- 
oughness remained as long as I knew him. Having _ gruntled bureaucrats and miserable weather. 

travelled with him, I could also recognize in that early After Harrow, Tom read Natural Sciences at Jesus 
diary his trademark dry and ironic humour, hidden in College, Cambridge. This academic subject captivat- 


620 


ed him and provided a continuum with his childhood 
interest in natural history. Owing to several circum- 
stances, the most prominent being boredom, he never 
took his degree. Instead, in the summer of 1932, at the 
age of twenty-one, he embarked on a serious journey, 
walking the length of the Atlantic coast of Norway 
on his own. Joined later by Reynold Bray, his close 
school friend from Harrow, the pair proceeded east- 
ward into Finland, making a winter trip by reindeer 
sledge through Lapland. In 1932, near the end of their 
trip, they trespassed into Russia. Tom told me they were 
arrested by villagers armed with pitchforks, who sus- 
pected them of spying. They subsequently spent three 
weeks in a Leningrad jail, being fed on increasingly 
weakened soup, until the British Consul secured their 
release. 

Exploration, travel and adventure were now Tom’s 
calling. From 1933-1935, he made a solitary expedi- 
tion to Southampton Island, northern Hudson Bay, to 
study a Snow Goose colony. He mapped the island 
under the auspices of the Royal Geographical Soci- 
ety and collected birds and mammals for the British 
Museum (Natural History). His longest journeys were 
made by dogsled in winter. From the Inuit of Coral Har- 
bor (on the island’s southern coast) he learned how to 
build an igloo, drive a dog team and hunt seal and 
caribou. When I once asked why he had an aversion to 
dumplings, he replied that during that first summer he 
had been given flour and baking powder by the Hud- 
son’s Bay Company Manager. Given no instructions 
and being too proud to ask what he should make, he 
existed on a tiresome diet of dumplings until rescued 
by the Inuit who showed him how to make bannock. 

Tom had no sooner returned to England in 1935 than 
he started to organize a larger expedition to South- 
ampton Island, and Foxe Basin. The five members of 
the British-Canadian Arctic Expedition, consisting of 
Tom, Graham Rowley, Reynold Bray, Pat Baird, Richard 
(Dick) Keeling and Peter Bennett (who later replaced 
Dick); arrived in Churchill in May of 1936. A smaller 
team consisting of Tom, Graham and Pat sailing in the 
Polecat, a 30-foot open whaleboat, established their 
main base at Coral Harbor. Each pursued his own 
investigation in biology, archaeology and geology. 
The expedition was later marred by the tragic death 
of Reynold Bray, who drowned after being blown out 
to sea in a collapsible boat. 

By the summer of 1938, Tom was alone in the field, 
at a camp in the Cape Dorset area of southern Baffin 
Island. He sent a message by Inuit dog team and radio 
to Montreal to Miss Ella Wallace Jackson (Jackie), a 
nurse from Nova Scotia whom he had met on the ship 
that took him back to England in 1935. He invited her 
to join him in the north. Jackie accepted, and the day 
after she arrived they were married, using a ring made 
from a brass fitting by the ship’s engineer. A few days 
later they headed north in a whaleboat. For the next 
eighteen months they conducted mapping and zoolog- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Tom Manning photograph by Borg Mesch, Kiruna, north- 
ern Sweden, 1932. 


ical studies on the west coast of Baffin Island. Jackie 
later published two books, /gloo for the Night, The 
University of Toronto Press, 1946 and A Summer on 
Hudson Bay, Hodder and Stoughton Ltd., 1949 about 
their travels together. Although Jackie and Tom spent 
many years together, the couple separated in the 1960s. 
Tom often visited her in Ottawa where she still resides. 

Late in 1939, Tom had a dream that caused him to 
head south; he and Jackie arrived in Cape Dorset on 2 
January 1940 to learn that World War II had been rag- 
ing for four months. Anxious to join the war effort, 
they discovered that the annual resupply ship — due 
in August — could not get them to Montreal before 
October. Tom decided they would make their own 
way back, continuing around the Foxe Basin coast of 
Baffin Island, hoping to reach Churchill by August. It 
was 20 January 1941, however, before they reached 
Churchill, after an epic journey of nearly 2500 miles 
by boat and dog team, mapping as they went. 

Tom received a commission in the Royal Canadian 
Navy Voluntary Reserve, but was soon seconded to 
the U.S. Army Engineers to provide advice on the sit- 


CARTER: THOMAS HENRY MANNING 


62) 


Tom Manning and Reynold Bray (1911-1936) on expedition 21 October 1932. Photograph T. Dahill of Kiruna, northern 
Sweden. 


| ing, construction and operation of an airfield and 
| weather stations on Southampton Island. He spent the 
| rest of the war on defence-related geodetic surveys in 
| the north, as well as the design and development of 
| Arctic clothing and equipment for the armed services. 


Tom served in the navy from 1941 to 1945, leaving 


| with the rank of Lieutenant Commander. 


Following the war, Tom’s services were sought by 
the Defence Research Board, the Geodetic Survey, the 


_ Geographical Branch, and other agencies of the Cana- 
_ dian Government. Assignments included the exploration 
_ of islands discovered in Foxe Basin by air photographs 


in 1948, a survey of Banks Island, an oceanographic 
survey in the Beaufort Sea, and many coastal surveys. 
Throughout his travels, Tom continued to collect spec- 


_imens of birds, mammals, plants and occasionally 
_ insects. The National Museum of Canada was one of 


the fortunate recipients of these valuable specimens. 


_ In 1949 Tom introduced the seventeen year old Andrew 


Macpherson to the Arctic. They became fast friends 
and made several expeditions together, collaborating 
on many reports of their findings. These journeys, 
with their combined achievements in the fields of sci- 
ence and exploration, are well known. Accounts of 


Tom’s work have appeared over the last few years in 
newspapers and journals around the world. In these 
articles writers have paid extensive tribute to his work. 
Many of his colleagues and acquaintances noticed that 
Tom was as much at ease at a formal dinner as on the 
snow-scoured rock of the Arctic Barrens. However, he 
preferred the familiar landscape of the Barrens and 
was able to make himself comfortable in surroundings 
and circumstances that many civilized people could 
not endure. 

After leaving the North, in 1941, Tom lived briefly 
in Montreal. Later, he and his wife bought a house in 
Ottawa on Linden Terrace where they entertained many 
of their northern friends. Tom started the Arctic Circle 
Club, 14 November 1947, when the guests no longer 
fitted around the huge oak dining-table that he had 
made himself. He was an expert craftsman, making 
cabinets for his house and building book shelves for his 
huge library. He and Jackie also spent hours patiently 
binding many of the collected volumes. 

When I first met him in 1965, Tom had given up his 
Arctic exploration and much of his scientific work. In 
the tradition of his grandfather T. A. Manning (TAM), 
he had become a farmer, an occupation which had 


622 


always interested him. However, Tom’s grandfather in 
Northampton, had forsaken farming and instead, found- 
ed a brewery continued by his son TEM. The prospect 
of this confining tradition was one of the forces that 
drove Tom to some of the most remote regions on the 
planet. However, he made some reconciliation with 
his heritage about 1960, by purchasing and working 
over 500 acres of hayfield, pasture and woodlot in the 
Merrickville countryside, some fifty km south of Otta- 
wa. He, and his Arctic friends, built a cabin on the largest 
portion of property which he also used for pasture. Tom 
went on to renovate a stone house and rework the fer- 
tile land into a self-sustaining farm with a huge and 
immaculate vegetable garden, cattle, sheep, chickens 
and geese. 

Tom and I talked a great deal about his northern trips. 
I finally convinced him that he should go back to his 
scientific work, and that I would make an excellent assis- 
tant! He cut back on the livestock and farming opera- 
tions, making them manageable for summer and occa- 
sional fieldwork. Our first trip was to western Canada 
collecting birds, mammals and plants. We travelled in 
a truck, which Tom had skillfully rebuilt as a home on 
wheels, complete with woodstove. He said it remind- 
ed him of caravan holidays as a child, and he always 
took immense pleasure in its rugged simplicity — often 
preferring to sleep in it, even when it was parked at 
home. 

My suggestion led to four decades of collaboration. 
I usually acted as assistant, and collected and prepared 
the birds with the hunting and skin preparation skills 
that Tom had taught me. There was ample opportunity 
for me to paint landscapes and wildlife studies on all 
the expeditions we made; and Tom was always encour- 
aging and supportive of my work. Occasionally I could 
provide help with illustrations or maps for his scientific 
papers. My first trip North with him was to Cape 
Henrietta-Maria, tagging polar bears for the Canadian 
Wildlife Service, under the direction of Dr. Charles 
Jonkel. Although I saw no bears, I fell in love with the 
land and, from then on, understood and shared Tom’s 
passion for this fabulous country. I was soon to see 
many bears, and we worked together from 1968 to 1973 
on the Circumpolar Polar Bear Project. 

During these travels, I had a firsthand chance to see 
Tom’s unique skills. As always, he collected as many 
types of specimens as he could, and kept extensive 
field notes and descriptions of everything we encoun- 
tered. Unlike many Arctic explorers of his time and 
previously, Tom had a broader range of interests, and 
documented everything he considered important. At 
the time, I did not fully comprehend either Tom’s skills 
or his decision-making process. Later, through field 
experience, I began to understand the brilliance of his 
mind and abilities. 

On one occasion he piloted an open boat in dense fog 
through seven miles (about 11 km) of the unforgiving 
waters of James Bay, using dead reckoning and expe- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


= 


Tom with beached hull of former 1949 expedition ship Nauya 
behind him, Igloolik, 1983, Photograph by Brenda 
Carter. 


rience to arrive at South Twin Island, exactly on target. 
On another occasion, attempting an open water cross- 
ing on the Belcher Islands, we ran into a dangerous 
pack-ice jam. This hazard was further compounded by 
an early snowstorm and a five-foot (1.5 m) tide rip. 
To stay meant being stranded. To leave meant facing 
rough water in an overloaded canoe. Tom’s choice lit- 
erally meant life or death. As usual, he made the right 
decision — elegant in its simple, straightforward truth. 
Despite the blinding snow he found a path through the 
heavy waves and got us safely ashore. He always kept 
part of his mind tuned to the essential core of any sit- 
uation, a trait which was the quiet binding of his life. 

From 1975 to 1986, Tom continued his collecting 
trips by truck in the summer, spending winters writing 
papers (a task he disliked) interspersed with his favour- 
ite seasonal activities of gardening, cutting fence-posts 
and filling the woodshed from his hardwood bush. 
Renewing his interest in his boyhood pursuit of eggs, 
he began to locate and weigh the eggs of different 
species. He could often be seen in nearby swamps with 
his odd, long-legged wooden table, a set of scales and 
an umbrella to shade the eggs. We both enjoyed the 
nest hunting and Tom became very interested in the 
results of his weight loss experiments, but the statistics 
proved daunting. I often wonder if a modern comput- 


2004 


er would have helped him with his puzzles. However, 
his resulting papers were successful, and the most re- 
| quested from scientists around the world. 

In 1983 Tom and I made our last trip north together. 
As we left Iqaluit for Igloolik, we had a poignant ex- 
change. Looking down at the shorelines and islands that 
| Tom had explored and mapped as a young man, | said 
that I thought I had come into his life fifteen years too 
late, and asked him what he felt as he looked down. He 
| said that he thought he also had arrived in the North 
fifteen years too late! At the age of seventy-five, he 
| continued to dream of an unmapped place and an era 
when the Arctic was truly beyond the reach of ordi- 
nary travel. 

On that trip, I noticed that Tom was walking badly; 
_ and for the first time since I met him, I realized that I 
| was able to outwalk him. This was startling and worry- 
| ing and we soon learned that he had Parkinson’s dis- 
| ease. However, he continued to work with great energy 
| on the farm. When Parkinson’s disease forced him to 
| give up his Arctic work, Tom donated his extensive col- 
lection of rare Arctic books along with his field journals 
and papers to the newly formed library in Iqaluit. He 
| also made a very generous gift of one million dollars 
| to the University of Cambridge towards their new lib- 


| rary at the Scott Polar Research Institute. He is remem- 
| bered there in a special room, the “Manning Archives”. 


Then, following a broken hip, Tom had to have an 


| operation. This trauma caused the Parkinson’s disease 
| to accelerate. After 10 years of his usual bravery and 
| independence, Tom became extremely ill. However, he 
| exhibited the same stoic patience that he had while 
| waiting out a blizzard. One of his last projects was to 
| slowly dig two new flower beds. His dry humour never 
| vanished, but the struggle became very difficult. In 
| November of 1998, at the age of 86 he died. He was in 
| hospital in Smiths Falls near his home, and remarkably 
| at peace. He left many legacies behind, including a 
| gift of $100 000 to the Ottawa Field Naturalist Club. 
| The council decided that 80% of interest from this 


bequest would go to assist authors with publication 
charges for the Canadian Field-Naturalist, primarily 


_ for northern papers. 


Tom was a renowned Arctic explorer and zoologist 


| whose life was characterized by a love of adventure, 
_ self-reliance, and an ever-inquiring mind. In fact as 
| Graham Rowley describes in his wonderful book, Cold 
Comfort: My Love Affair with the Arctic, (McGill-— 


Queen’s University Press, 1996) what Tom accom- 


_ plished will never be equalled. He was also a wonder- 


ful companion and brilliant teacher, showing by exam- 
ple how to use intuition and logic to unravel a crisis and 
how to see both facts and beauty in the natural world. 


| He was quiet to the point of being taciturn, and slow to 
| join in the mindless customs of society. But, his loy- 


alty to his friends and to his family in England was 
complete. Those who were privileged to know him well 
miss him in many, many ways. 


CARTER: THOMAS HENRY MANNING 


623 


Other tributes to Tom have appeared in several news- 
paper articles as well as an obituary in Arctic 52 (1): 
104-105 [March 1999] by Andrew H. Macpherson. 
Tributes to his disciple Andrew Hall Macpherson (1932- 
2002) and Manning’s role as his mentor have appeared 
in The Daily Telegraph (London, England) 22 June 
2002 and in Arctic 55(4): 403-406 (December 2002) 
by Frank L. Miller. 


Awards 

1944 — Bruce Medal of the Scottish Geographical Society 
and the Philosophical Society of Edinburgh. 

1948 — Patron’s Medal of the Royal Geographical Society 

1959 — Guggenheim Fellowship held for Arctic research 

1974 — Officer of the Order of Canada 

1977 — Massey Medal of the Royal Canadian Geographical 
Society 

1977 — Queen’s Jubilee Award 

1979 — Honorary LLD from McMaster University 

1992 —Doris Huestis Speirs Award in recognition of his 
pioneering work on birds in the Canadian North 


Positions in Scientific and Professional Societies 

Fellow: Royal Geographical Society; Royal Canadian Geo- 
graphical Society (also a term as Director) Arctic 
Institute of North America (also Executive Director 
1955-1956). 

Member: American Ornithological Union; Wilson Ornitho- 
logical Club; Cooper Ornithological Club; Wildlife 
Society; American Society of Mammalogists; Society 
for American Archaeology. Founder: Arctic Circle 
Club; 1947 (also Secretary 1948-1950) 

Research Associate: Canadian Museum of Nature (1970-1990) 


Bibliography 

Publications 

Manning, T. H. 1936. Some notes on Southampton Island. 

Geographical Journal 88: 232-242. 

Manning, T. H. 1942. Remarks on the physiography, Eskimo 

and mammals of Southampton Island. Canadian Geograph- 

ical Journal 25: 17-35. 

Manning, T. H. 1942. Notes on some fish of the eastern 

Canadian Arctic. Canadian Field-Naturalist 56: 128-129. 

Manning, T. H. 1942. Blue and Lesser Snow geese on South- 

ampton and Baffin Islands. Auk 59: 158-175. 

Manning, T. H. 1943. Notes on the coastal district of the east- 

ern barren grounds and Melville Peninsula from Igloolik to 

Caper Fullerton. Canadian Geographical Journal 26: 84-105. 

Manning, T. H. 1943. The Foxe Basin coasts of Baffin Island. 

Geographical Journal 101(5, 6): 225-251. 

Manning, T. H. 1943. Notes on the mammals of south and 

central west Baffin Island. Journal of Mammalogy 24: 

47-59. 

Manning, T. H. 1943. Notes on the birds of Southampton 
Island, Baffin Island and Melville Peninsula. Auk 60: 504- 
536. [prepared partially from the notes of R. J. O. Bray, 
who died on the expedition] 

Manning, T. H. 1944. Hunting implements and methods of 
the present day Eskimos of north-west Hudson Bay, Mel- 
ville Peninsula, and south-west Baffin Island. Geographi- 
cal Journal 103: 137-152. 

Manning, T. H., and E. W. Manning. 1944. The preparation 
of skins and clothing in the eastern Canadian Arctic. Polar 
Record 28: 156-169. 


624 


Manning, T. H. 1944. Lesser Snow and Blue Geese on South- 
ampton Island. Auk 61: 146-147. 

Manning, T. H. 1946. Ruins of Eskimo stone houses on the 
east side of Hudson Bay. American Antiquity 11(3) 201- 
202. 

Manning, T. H. 1946. Bird and mammal notes from the east 
side of Hudson Bay. Canadian Field-Naturalist 60: 71-85. 

Manning, T. H. 1947. Explorations on the east coast of Hud- 
son Bay. Geographical Journal 109: 58-75. 

Manning, T. H. 1948. Eskimo stone house ruins on the east 
side of Hudson Bay: A correction. American Antiquity 13 
(3). 

Manning, T. H. 1948. Pipestems of the Caribou Eskimos. 
American Anthropologist 50: 162-163. 

Manning, T. H. 1948. Notes on the country, birds and mam- 

mals west of Hudson Bay between Reindeer and Baker 

lakes. Canadian Field-Naturalist 62: 1-28. 

Manning, T. H. 1949. The birds of northwestern Ungava. 

Pages 155-224 in A Summer on Hudson Bay by Mrs. T. 

H. Manning. Hodden and Stoughton Ltd. 

Manning, T. H. 1949. The varying lemming in captivity in 
Ottawa. Arctic Circular 3: 20-21. 

Manning, T. H. 1950. Tidal observations in Arctic waters: 
Notes on the tides along the south Hudson Bay and west 
James Bay coasts. Arctic 3: 95-100. 

Manning, T H. 1950. Eskimo stone houses in Foxe Basin. 

Arctic 3: 108-112. 

Manning, T. H. 1950. A mixed Cape Dorset-Thule site on 

Smith Island, east Hudson Bay. Annual Report of the 

National Museum of Canada 1949-50, Bulletin 123: 64-71. 

Manning, T. H., et al. 1950. The voyage of C. G. M. V. Nauja 

to Foxe Basin in 1949. 158 pages, plus maps, photographs. 

Manning, T. H. 1951. Remarks on the tides and driftwood 
strand lines along the east coast of James Bay. Arctic 4: 
122-130. 

Manning, T. H. 1952. Birds of the west James Bay and south- 
erm Hudson Bay coasts. National Museum of Canada Bul- 
letin 125: 1-114. 

Manning, T. H., and D. F. Coats. 1952. Notes on the birds 

of some James Bay islands. Annual Report of the National 

Museum Canada 1950-51, Bulletin 126: 195-207. 

Manning, T. H., and A. H. Macpherson. 1952. Birds of the 

east James Bay coast between Long Point and Cape 

Jones. Canadian Field-Naturalist 66: 1-35. 

Manning, T. H. 1953. Narrative of an unsuccessful attempt 

to circumnavigate Banks Island by canoe in 1952. Arctic 

6: 170-197. 

Manning, T. H. 1953. Notes on the fish of Banks Island. 

Arctic 6: 276-277. 

Manning, T. H. 1954. Remarks on the reproduction, sex ratio, 

and life expectancy of the varying lemming, Dicrostonyx 

groenlandicus, in nature and captivity. Arctic 7: 36-48. 

Manning, T. H. 1956. The northern red-backed mouse, Cleth- 

rionomys rutilus (Pallas), in Canada. National Museum of 

Canada Bulletin 144: 1-67. 

Manning, T. H. 1956. Narrative of a second Defence Research 

Board expedition to Banks Island, with notes on the coun- 

try and its history. Arctic 9(1, 2): 1-77. 

Manning, T. H., E. Q. Hohn, and A. H. Macpherson. 1956. 

The birds of Banks Island. National Museum of Canada 

Bulletin 143: 1-144. 

Manning, T. H., and A. H. Macpherson. 1958. The mammals 
of Banks Island. Arctic Institute of North America Tech- 
nical Paper 2: | -74. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Macpherson, A. H., and T. H. Manning. 1959. The birds and 
mammals of Adelaide Peninsula, N.W.T. National Museum 
of Canada Bulletin 161: 1-63. 

Manning, T. H. 1960. The relationship of the Peary and bar- 
ren ground caribou. Arctic Institute of North America Tech- 
nical Paper 4: 1-52. 

Manning, T. H. 1961. Comments on “Carnivorous Walrus 
and Some Arctic Zoonoses”. Arctic 14(1): 76-77. 

Manning, T. H., and A. H. Macpherson. 1961. A biological 
investigation of Prince of Wales Island, N.W.T. Transactions 
of the Royal Canadian Institute 33: 116-239. 

Macpherson, A. H., and T. H. Manning 1961. Pack dogs in 

the Canadian Arctic. Polar Record 10: 509. 

Manning, T. H. 1964. Age determination in the polar bear, 

Ursus maritimus Phipps. Canadian Wildlife Service Occa- 

sional Paper 5: 1-12. 

Manning, T. H. 1964 Geographical and Sexual Variation in 

the Long-tailed Jaeger, Stercorarius longicaudus Vieillot. 

Biological Papers of the University of Alaska. (7): 1-16. 

Manning, T. H. 1969 Wreckage on Banks Island. Beaver 

Autumn: 21. 

Manning, T. H. 1971. Geographical variation in the polar bear 

Ursus maritimus Phipps. Canadian Wildlife Service Report 

Series 13: 1-26. 

Manning ,T. H. 1974. Variations in the skull of the bearded 

seal. Biological Papers of the University of Alaska (16). 

Manning, T. H. 1976. Birds and mammals of the Belcher, 
Sleeper, Ottawa and King George islands, Northwest Ter- 
ritories. Canadian Wildlife Service Occasional Papers 28: 
1-42, maps, tables. 

Morrison, R. I. G., T. H. Manning, and J. A. Hager. 1976. 
Breeding of the Marbled Godwit, Limosa Fedoa, in James 
Bay. Canadian Field-Naturalist 90: 487-490. 

Morrison, R. I. G., and T. H. Manning 1976. First breed- 
ing records of Wilson’s Phalarope for James Bay, Ontario. 
Auk 93(3): 656-657. 

Manning, T. H., and B. Carter. 1977. Incidence of runt eggs 
in the Canada Goose and Semipalmated Sandpiper. Wilson 
Bulletin 89: 469. 

Manning, T. H. 1978. Measurements and weights of eggs 
of the Canada Goose, Branta canadensis, analyzed and 
compared with those of other species. Canadian Journal 
of Zoology 56: 676-687. 

Manning, T. H. 1979. Density and volume corrections of 

eggs of seven passerine birds. Auk 96: 207-211. 

Manning, T. H. 1981. Analysis of weight lost by eggs of ele- 

ven species of birds during incubation. Canadian Field- 

Naturalist 95: 63-68. 

Manning, T. H. 1981 Birds of the Twin Islands, James Bay, 

N.W.T., Canada. National Museum of Natural Sciences, 

Syllogeus 30: 1-50. 

Manning, T. H. 1982. Daily Measurements of Variation in 
weight loss of eggs of seven Passerine Species before and 
during natural incubation. National Research Council of 
Canada 60: 3143-3149. 

Barker, A. J., J. L. Eger, R. L. Peterson, and T. H. Manning. 
1983. Geographic variations and taxonomy of Arctic 
Hares. Acta Zoologica Fennica 174: 45-48. 


Unpublished Reports 

Manning, T. H. 1944. Descriptive report to the Geodetic 
Service of Canada on a survey of the east Hudson Bay 
coast. 64 pages plus maps, photographs. 


2004 


' Manning, T. H. 1945. Report on Exercise Lemming to the 
Geodetic Service of Canada. 21 pages plus 4 pages, plus 
photographs. 

|Manning, T. H. 1945. Descriptive report to the Geodetic 

Service of Canada on a survey of northern Manitoba and 
southeast Keewatin. 54 pages, plus photographs. 

Manning, T. H. 1945. Technical report to the Geodetic Ser- 

vice of Canada on a survey of northern Manitoba and 
southeast Keewatin. 179 pages, plus maps, photographs. 

Manning, T. H. 1946. Report to the Geodetic Service of 

Canada on Exercise Muskox and on the Cambridge Bay 

___ observation station. 25 pages, plus photographs. 

| Manning, T. H. 1946. Descriptive report to the Geodetic 

Service of Canada on a survey of Ungava, east Hudson 

Bay, and the south coast of Hudson Strait. 133 plus 49 pages, 

photographs and maps. 

| Manning, T. H. 1946. Technical report to the Geodetic Ser- 

vice of Canada on a survey of Ungava, east Hudson Bay, 

| and the south coast of Hudson Strait. 90 pages, maps. 

| Manning, T. H. 1947. Preliminary report on a background 

| study of the caribou, Rangifer caribou caribou (Gmelin) and 

Rangifer arcticus caboti Allen of the Labrador Peninsula 

and the Province of Quebec north of the St. Lawrence. 

(Mimeo). 

| Manning, T. H. 1947. Descriptive report to the Geodetic Ser- 

vice of Canada on a survey of the west James Bay and 

southern Hudson Bay coasts. 166 pages, plus maps, pho- 
tographs. 

| Manning, T. H. 1947. Technical report to the Geodetic Ser- 

vice of Canada on a survey of the west James Bay and south- 

ern Hudson Bay coasts. 64 pages, plus maps. 

| Manning, T. H. 1948-49. Reports to Defence Research Board. 

1. South coast of Hudson Bay, 147 pages, plus photographs, 

maps, charts, etc. II. West side of James Bay, 185 pages, plus 

photographs, etc. FV. East coast of James Bay, 295 pages, 
plus photographs, etc. 


Professional Experience 

| 1931 (summer) — Traveled in Iceland and Faro Islands. 

1932-33 (winter) — Traveled through Norway, Sweden, Fin- 

land, and Russia (Bergen to Murmansk) on foot and 
with reindeer. 

| 1933-35 — Mapped Southampton Island and also studied 

and made collections of birds and mammals. 
1936-41 — Leader, surveyor and ornithologist on the British 

Canadian — Arctic Expedition, working in Southamp- 

ton and Baffin islands and Repulse Bay areas: includ- 

ed two years in Foxe Basin which was circumnavi- 

| gated by whale boat. 

1941-45 — Royal Canadian Navy. Seconded as an advisor to 

the U.S. Army in connection with the sighting and con- 

struction of the air field and establishment by tractor 

train of outlying weather stations on Southampton 

| Island; and later to the Geodetic Service (Canada) to 

| fix ground controls for air photographic survey of 

northern Canada (until 1947). 

| 1944 — Geodetic work in Ungava by aircraft and Peterhead 

| boat. 

j 1945 — Geodetic work in west Hudson Bay by aircraft. 

| 1946 — Geodetic work in Ungava, east Hudson Bay and west 

Hudson Strait by aircraft and Peterhead boat. 
_ 1947 — Geodetic work by canoe along the west James Bay 
and southern Hudson Bay coasts. 


CARTER: THOMAS HENRY MANNING 


625 


1948 — Consultant to Defence Research Board. 

1949 — Leader, Geographical Bureau’s expedition in C. G. 
M. V. Nauja to “new” islands in Foxe Basin: includ- 
ed circumnavigation of Foxe Basin. Soundings and 
tidal observations taken. 

1950 — Canoe journey to collect birds and mammals along 
east James Bay. 

1951 — Leader, Defence Research Board expedition to the 
Beaufort Sea and Master C.G.M.V Cancolim (includ- 
ed “circumnavigation” of Alaska). About 4000 miles 
of soundings taken. 

1952 — Leader, Defence Research Board expedition to Banks 
Island: included attempted circumnavigation of the 
Island by canoe. 

1953 — Completion of circumnavigation of Banks Island. 

1954 — Worked up results of Banks Island Expedition under 
contract with Defence Research Board, and others. 

1960 (June to July) — West Coast of North America with 
Diana Rowley 

1967 — Introduction of caribou from Coats Island to South- 
ampton Island for Canadian Wildlife Service. 

1968 (June 14 — August) — Western Canada collecting by 
truck assisted by Brenda and Charlotte Carter; (Sep- 
tember) — Cape Henrietta-Maria with Brenda Carter, 
Dick Russell and Chuck Jonkel for Canadian Wildlife 
Service. 

1969 — Alaska collecting by truck assisted by Z. E. Ellshoff 

1970 (July 21 — August) — South Twin Islands, polar bear 
trapping with Brenda Carter for Canadian Wildlife 
Service. : 

1971 (May 2 — September) — Belcher Islands, Sleeper Islands 
and Ottawa Islands with Brenda Carter for Canadian 
Wildlife Service. 

1972 (May — June) — UK and Norway measuring bear 
skulls with Charlotte Carter; (September 10 — Octo- 
ber 18) -North Twin Island, polar bear trapping with 
Brenda Carter for Canadian Wildlife Service. 

1973 (March 22 — April 17) — Fort George and Belcher 
Islands, helicopter polar bear tagging with Brenda 
Carter for Canadian Wildlife Service; (May 2 — Au- 
gust 1) — North Twin Island, polar bear work with 
Brenda Carter for Canadian Wildlife Service; (August 
13 — September 12) — Foxe Basin helicopter work 
with Brenda Carter for Canadian Wildlife Service. 

1975 (May 15 — July 16) — Collecting at North Point, James 
Bay. 

1977 — Yukon collecting by truck and making a floral and 
faunal inventory of the Dempster Highway assisted 
by Jan Rowell and Wendy Earl (who departed Daw- 
son City). Robert Porsild and his wife were visited. 

1980 — New Brunswick collecting by truck. 

1982 — Manitoulin Island collecting by truck assisted by Doug 
Perkins 

1983 — Igloolic Wildlife description. Advisor to adult Inuit 
for Igloolic Education Council. 

1983 — USA (Mississippi, Louisiana, Nevada, Utah and 
Texas) collecting by truck assited by S. C. Tuutz. 

1986 — Yukon collecting by truck (age 77) assisted by Brid- 
get Ryan. 

Most of the specimens collected on these trips were sold either 
to the National Museums of Canada or Carleton 
University. 


A Tribute to Victor Kent Prest 1913-2003 


A. S. DyKE!. D. A. Hopcson!, E. L. BouSFIELD?, and R. E. BEDFORD? 


: Terrain Sciences Division, Geological Survey of Canada, 601 Booth Street, Ottawa, Ontario K1A OE8 Canada 
~ 301-540 Dallas Road, Victoria, British Columbia V8V 4X9 Canada 


; 524 Penhill Avenue, Ottawa, Ontario K1G 0V6 Canada 


Dyke, A. S., D. A. Hodgson, E. L. Bousfield, and R. E. Bedford. 2004. A Tribute to Victor Kent Prest 1913-2003. Canadian 


Field-Naturalist 118(4): 626-629. 


With the death of Dr. Victor Kent Prest on 2003 Sep- 
tember 26, Canada lost one of its most distinguished, 
best-known, and most influential Quaternary geolo- 
gists. Vic was active as a leader of Canadian Quater- 
nary science for a remarkably long time and he posi- 
tively influenced the careers of almost everyone else 
in the field in this country as well as his many friends 
abroad. His work became basic to discoveries by other 
scientists in related fields such as plant and animal 
biogeography, both terrestrial and aquatic*>°. He came 
to personify the Quaternary Geology of Canada both 
domestically and internationally. This identity arose 
from his unstinting and deep interest in the subject, 
his long view of the evolution of ideas, his intimate 
knowledge of the ever-growing knowledge base, and 
especially the very genuine interest he took in the work 
of all others. Tentative, junior researchers and graduate 
students always met with as much respect and encour- 
agement as did his senior colleagues and Vic often took 
the time to send notes of encouragement and compli- 
ment. 

Victor K. Prest was born in Edmonton Alberta, 2 
April 1913, the only child of John (Jack) Prest and 
Elizabeth Buckley (Prest). His early education was 
obtained at the MacKay Avenue School in Edmonton. 
In 1925 the family moved to Toronto for a brief period, 
and relocated in London, Ontario, where Vic completed 
his early schooling at Tecumseh Public School and 
London South Collegiate High School. In 1930 the 
Prests moved to Winnipeg where Victor completed his 
secondary education at Kelvin High School. Unreal- 
ized by Vic, his future wife Patricia Horder was also 
attending Kelvin, in the same grade, but in a different 
classroom. Her father, a Canadian Pacific Railway offi- 
cial, had been transferred from Montreal to Winnipeg 
in 1926. Vic received his B. Sc. (Honours) from the 
University of Manitoba in 1935 and his M. Sc. a year 
later. Growing stronger during these university days 
was his friendship with Patricia Horder who was active 
in music, drama, and art, and in the Canadian Girl 
Guide Association. 

Vic worked as a summer assistant for the Manito- 
ba Mines Branch in 1934 and the Geological survey 
of Canada in 1935 and 1936. While registered for 
Ph.D. studies at the University of Toronto, Vic was 


Party Chief, Ontario Department of Mines, from 1937- 
1940, mainly mapping bedrock outcrops in Northern 
Ontario. In 1941 Vic received his Ph.D. degree at “Tor- 
onto” where he also served as an assistant in both the 
Geology Department and the Royal Ontario Museum 
and later as Lecturer. The newly minted Dr. Prest 
joined the International Nickel Company as a geolo- 
gist for a year (1941-1942). 

Vic and Pat were married in Toronto in 1942 and 
moved immediately to Harbour Grace, Newfoundland, 
where Vic served with the RCNVR from 1942-1945. 
Their first child, Sherron (Sherry) Gail was born there 
in 1944. Following the surrender of Nazi Germany in 
early May, 1945, Lieutenant Prest was a member of the 
naval party accepting the surrender of submarine U190 
in St. John’s Harbour. 

During the next five years Vic was a permanent staff 
member of the Ontario Department of Mines, based in 
Toronto, where his son Wayne Horder Prest was born 
in 1946. In 1950, he transferred to the Geological Sur- 
vey of Canada in Ottawa. Vic served as Chief of the 
Pleistocene Engineering and Groundwater Section, 
first in a third-floor office of the Victoria Memorial 
Museum building and, following its completion in 
1960, in the newly constructed Geological Survey 
building on Booth Street. Vic formally retired from 
government service in 1979, but maintained a con- 
sultantship office at the Booth St. building until very 
recently. 

Dr. Victor K. Prest is probably best known because 
of the Glacial Map of Canada (Prest et al. 1968), a 
work of such veracity that it has yet to be superceded 
for the country as a whole. This map followed an ear- 
lier version initiated by J. Tuzo Wilson, to which Vic 
contributed as co-compiler (Wilson et al. 1959). The 
1968 Glacial Map was followed by the beautiful blue- 
shaded “pancake map” depicting the Retreat of Wis- 
consin and Recent Ice in North America (Prest 1969), 
and his masterful Quaternary Geology chapter in the 
Geology of Canada volume (Prest 1970). That trilogy 
was the masterpiece of a generation, which adorned 
many an academic hallway and classroom. Its impact 
places it among the most outstanding contributions of 
the Geological Survey of Canada (GSC) and fixes it as 
its premier Quaternary contribution. It was the most 


626 


2004 


DYKE, HODGSON, BOUSFIELD, and BEDFORD: VICTOR KENT PREST 


627 


VICTOR KENT PREST 


cogent, persuasive, and attractively illustrated synthesis 
available and it has served as the baseline for almost 
everything else that has followed. 

As a great builder of Quaternary science in Canada, 
Vic had wide knowledge of geological and other sci- 


_ ences. He resumed mapping the Precambrian geology 


of northern Ontario for ODM (1945-1950) where he 
authored more than 30 maps and annual reports. He 
was one of the first to combine Quaternary and bedrock 
mapping with the release of his map of Red Lake- 
Lansdowne House. His first Quaternary publication was 
the Pleistocene geology of the Vermillion River system 
with special reference to placer gold (Prest 1949). 
Vic’s early Pleistocene colleagues at GSC includ- 
ed Jack Armstrong, Wes Blake, Jr., Bruce Craig, Bob 


Fulton, John Fyles, Nelson Gadd, Eric Henderson, 
Owen Hughes, Bert Lee, and Archie Stalker, all of 
whom were to become major regional authorities, and 
Jaan Terasmae and Bob Mott, leaders in the develop- 
ment of Quaternary paleoecology in Canada. Addi- 
tion of his academic and provincial geological survey 
friends, Bill Matthews (University of British Colum- 
bia), Earl Christiansen (Saskatchewan), Alexis Drei- 
manis (University of Western Ontario), Con Gravenor 
(University of Windsor), Paul Karrow (Waterloo Uni- 
versity), Pierre LaSalle (Quebec), and John Elson 
(McGill University) largely completes the Prest cohort 
of Canadian Quaternary geologists. 

Although he departed senior management for full- 
time research in 1964, even those who joined the Sur- 


628 


vey later, and many in outside agencies, identified the 
Quaternary group with Vic Prest more than with any- 
one else. Vic became a sort of mentor-at-large, with 
great moral authority and prestige, and engendered 
pride in the work of others. By building Quaternary 
science at the Survey and by mentoring and infor- 
mally supervising graduate students, Vic profoundly 
contributed to the Canadian Quaternary community. 

Vic was an enthusiastic participant in numerous field 
excursions in southern Canada, the northern USA, 
Alaska, and internationally. His GSC fieldwork took 
him on a High Arctic voyage that reached Ellesmere 
Island (Prest 1958), mapping projects in Ontario (Prest 
1963), Quebec (Prest 1966; Prest and Hode-Keyser 
1977) and the Maritimes. He had a special passion for 
the Maritimes, where his map of Prince Edward Island 
(Prest 1973) remains seminal, and where, with his pro- 
tégé Douglas Grant, he formulated the concept of an 
Appalachian Ice Complex (Prest and Grant 1969; Prest 
et al. 1972; Prest 1977) to encapsulate the style of 
regional glaciation, as distinct from Laurentide glacia- 
tion. This model is still favoured by regional geolo- 
gists. The Magdalen Islands in the Gulf of St. Law- 
rence long held his interest, because of the perplexing 
lack of direct evidence of Late Wisconsin glaciation 
and its record of older events (Prest et al. 1976). This 
evidence led him to portray on his ice recession map 
what Doug Grant termed “Prest’s Problematical Pleis- 
tocene Prong”, a reference to the glaciological improb- 
ability of leaving the Magdalen Islands ice free while 
allowing ice to extend to the edge of the Continental 
Shelf. The portrayal revealed Vic’s preference for 
field evidence over theory. 

Vic formally retired from GSC in 1978, and formed 
Veekay Consultants (1978-1994) to receive Ontario 
Geological Survey (OGS) surficial mapping contracts 
in northwestern Ontario and a Department of Indian 
Affairs and Northern Development contract to work 
in the Bebensee Lake area (Prest 1985). He continued 
fieldwork into his seventies, including long traverses 
into dense bush around the famous Red Lake gold 
mining camp. This work yielded a series of detailed 
maps (e.g., Prest 1980) that formed the basis for recent 
drift prospecting programs. His major syntheses includ- 
ed Canada’s Heritage of Glacial Features (Prest 1983), 
the Late Wisconsin Glacier Complex paper and map 
(Prest 1984), a synthesis of glacial lake history in the 
Lake Ontario basin with Ernie Muller (Muller and 
Prest 1985), and maps of North American paleogeog- 
raphy (Dyke and Prest 1987). His two final publications 
dealt with the history of interpretation of the Laurentide 
Ice Sheet (Prest 1990) and a more specific passion, the 
long-distance dispersal of the distinctive erratics that 
he termed “omarsi” from the central part of the ice sheet 
(Prest et al. 2000). 

Vic was an accomplished athlete and maintained a 
life-long interest in several sports. During the half cen- 
tury of his residence in Ottawa, Vic’s love of tennis 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


became super-seded by a passionate devotion to curl- 
ing. He was a charter member of the City View Curl- 
ing Club, founded in 1957, and President the follow- 
ing year. Over the years, he won many curling bonspiels 
both in men’s and mixed rink events, and was ever a 
formidable shot-maker and curling strategist. He was 
Honorary Governor General’s Curling Club President 
(1972), Canadian Branch (Royal Caledonia Curling 
Club), Council of Management (1978-1980) and Life 
Member (1986). With support from the Ottawa Val- 
ley Curling Association, Vic introduced Junior Curl- 
ing in Eastern Ontario in 1975, and co-ordinated play 
with the Canadian Branch in Quebec. He also intro- 
duced “Little Rock” curling in 1983, and organized 
Junior and Little Rock bonspiels for the OVCA over 
a period of ten years. After more than 50 continuous 
years of winter curling, he “pushed” his last rock at 
age 89. 

Vic’s professional career accomplishments received 
due recognition in the form of the Johnston Medal 
from the Canadian Quaternary Association (the first 
awarded), the Gold Medal from the Royal Canadian 
Geographical Society, the Kirk Bryan and the Distin- 
guished Career Awards from the Quaternary Geology 
and Geomorphology Division of the Geological Society 
of America (1993), election as a Fellow to join the 
distinguished scientists of the Royal Society of Cana- 
da, and election as Honorary Member of the Interna- 
tional Union for Quaternary Research. 

Vic and Pat enjoyed more than sixty years of mutual 
accomplishment. He loved the out-of-doors and experi- 
enced great pleasure whenever he had a chance to stand 
on a good piece of Precambrian bedrock. Vic was also 
very generous in his support of charitable organiza- 
tions, and agencies promoting wildlife conservation 
and the natural sciences. He will be greatly missed by 
his family, and by his many friends and scientific col- 
leagues. 


References 

Dyke, A. S., and V. K. Prest. 1987. The Late Wisconsin and 
Holocene history of the Laurentide Ice Sheet. Géogra- 
phie physique et Quaternaire 41: 247-264. 

Muller, E. H., and V. K. Prest. 1985. Glacial lakes in the 
Ontario basin. Edited by P. F. Karrow, and P. E. Calkin in 
Quaternary Evolution of the Great Lakes. Geological 
Association of Canada, Special Paper 30: 213-229. 

Prest, V. K. 1939. Geology of the Keezhik-Miminiska Lake 
area. Ontario Department of Mines, Annual Report 48, 
Part 6, 8 pages. 

Prest, V. K. 1941. The Precambrian of the Miminiska-Port 
Hope area. Unpublished Ph.D. thesis, University of Toronto, 
93 pages. 

Prest, V. K. 1949. The Pleistocene geology of the Vermillion 
River system near Capreol, District of Sudbury, Ontario 
(with special reference to placer gold), Ontario Department 
of Mines Open File. 

Prest, V. K. 1952. Notes on the geology of parts of Ellesmere 
and Devon islands. Geological Survey of Canada, Paper 
52-32, 15 pages. 


2004 


| Prest, V. K. 1963. The Red Lake-Lansdowne House area, 
northwestern Ontario — Surficial geology. Geological Sur- 
vey of Canada, Paper 63-6, 23 pages. 

Prest, V. K. 1966. Glacial studies, northeastern Ontario and 
northwestern Quebec. In Report of Activities May to Octo- 
ber 1965. Geological Survey of Canada, Paper 66-1: 202- 
203. 

Prest, V. K. 1969. Retreat of Wisconsin and Recent ice in 
North America. Geological Survey of Canada, Map 1257A, 
scale 1:5 000 000. 

Prest, V. K. 1970. Quaternary geology of Canada. Edited by 
R. J. W. Douglas in Geology and Economic Minerals of 
Canada. Economic Geology Series (1): 676-764. 

Prest, V. K. 1973. Surficial deposits of Prince Edward 
Island. Geological Survey of Canada, Map 1366A, scale 
1:253 444. 

Prest, V. K. 1977. General stratigraphic framework of the Qua- 
ternary in eastern Canada. Géographie physique et Qua- 
ternaire 31: 7-14. 

| Prest, V. K. 1980. Quaternary geology of the Red Lake area, 
District of Kenora. Ontario Geological Survey, Preliminary 
Map 2484, scale 1:50 000. 

Prest, V. K. 1983. Canada’s heritage of glacial features. Geo- 
logical Survey of Canada, Miscellaneous Report 28: 119 
pages. 

Prest, V. K. 1984. Late Wisconsinan Glacier Complex. Edited 
by R. J. Fulton, in Quaternary Stratigraphy of Canada — 
A Canadian contribution to IGCP Project 24. Geological 
Survey of Canada, Paper 84-10: 21-38. 

Prest, V. K. 1984. Late Wisconsinan Glacier Complex. Geo- 
| logical Survey of Canada, Map 1584A, 1:7 500 000. 
Prest, V. K. 1985. Glacial geology of the Bebensee Lake map- 

area (NTS 86M), Great Bear Lake region, Northwest Ter- 
ritories. Edited by J. A. Brophy in Contributions to the 
Geology of the Northwest Territories, 63-70. 

Prest, V. K. 1990. Laurentide ice-flow patterns: a historical 
review, and implications of the dispersal of Belcher Islands 
erratics. Géographie physique et Quaternaire 44: 113-136. 

Prest, V. K., and J. A. Donaldson. 1963. Bedrock-drift dis- 
tribution, Red Lake-Lansdowne House area, Ontario. Geo- 
logical Survey of Canada, Map 4-1963. 

Prest, V. K., J. A. Donaldson, and H. D. Moores. 2000. 
The Omar story: The role of Omars in assessing glacial his- 


DYKE, HODGSON, BOUSFIELD, and BEDFORD: VICTOR KENT PRES1 


tory of west-central North America. Géographie physique 
et Quaternaire 54: 257-270. 

Prest, V. K., and D. R. Grant. 1969. Retreat of the last ice 
sheet from the Maritime Provinces — Gulf of St. Lawrence 
region. Geological Survey of Canada, Paper 69-33, 15 pages. 

Prest, V. K., D. R. Grant, H. W. Borns, I. A. Brookes, R. H. 
MacNeill, J. G. Ogden, J. F. Jones, T. W. Hennigar, and 
M. L. Parsons. 1972. Quaternary geology, geo-morphology 
and hydrogeology of the Atlantic Provinces. XXIV Inter- 
national Geological Congress, Excursion Guidebook A61- 
C61, 79 pages. 

Prest, V. K., D. R. Grant, and V. Rampton. 1968. Glacial 
Map of Canada. Geological Survey of Canada, Map 1253A, 
scale 1:5 000 000. 

Prest, V. K., and J. Hode-Keyser. 1977. Geology and engi- 
neering characteristics of surficial deposits, Montreal Island 
and vicinity, Quebec. Geological Survey of Canada, Paper 
75-29, 29 pages. 

Prest, V. K., J. Terasmae, J. V. Matthews, and S. Lichti- 
Federovich. 1976. Late-Quaternary history of Magdalen 
Islands, Quebec. Maritime Sediments, 12(2): 39-59. 

Wilson, J. T., G. Falconer, W. H. Mathews, and V. K. 
Prest. 1959. Glacial Map of Canada. Geological Associ- 
ation of Canada, scale 1:3 801 600. 


Examples of research publications in periph- 
eral fields of science for which the work of V. 
K. Prest was basic: 

Bousfield, E. L., and M. L. H. Thomas 1975. Postglacial 
changes in distribution of littoral marine invertebrates in 
the Canadian Atlantic Region. Proceedings of the Nova Sco- 
tia Institute of Science 27, Supplement 3: 37-42, 8 figures. 

Dadswell, M. J. 1974. Distribution, ecology, and postglacial 
dispersal of certain crustaceans and fishes in eastern North 
America. National Museum of Canada Publications in 
Zoology 11: 110 pages, 20 maps. 

Pielou, E. C. 1991. After the Ice age. University Chicago Press. 
Chicago. 366 pages. 


Acknowledgments 

For providing biographical material, photographs, and per- 
sonal recollections, the authors are especially grateful to 
Patricia Prest (wife) and Sherry Armstrong (daughter). 


Book Reviews 


ZOOLOGY 


Birds of Australia (Seventh edition) 


By Ken Simpson and Nicolas Day. 2004. Princeton Univer- 
sity Press, 41 William Street, Princeton, New Jersey, 
USA, 08540-5237. iv + 92 pages, U.S.$39.50. 


When Simpson and Day’s first edition appeared (as 
Birds of Australia, later changed to The Princeton Field 
Guide to the Birds of Australia) it meant Australia had 
a bird guide that was equivalent to North America’s 
National Geographic guide (Dunn 2002). It provided 
complete coverage of Australian birds in color, with 
accompanying text and range maps. Curiously the print- 
ing was, and still is, poorer than that of the National 
Geographic. 

As with the original this book covers all six states 
(including Tasmania), three territories and 10 island 
territories. The new edition has a double page map of 
the continent, but does not show the island territories. 

This revised edition has undergone significant 
changes from the original. Although much of the con- 
tent is the same it has been reorganized. More impor- 
tant 19 of the plates have been repainted (actually I 
counted 24 plates with changes). The illustrations now 
include more plumage variations than before, such as 
female, juvenile, winter and different races. In addition 
to over 2000 colour images there are almost 1000 sup- 
plementary black-and-white drawings. One noticeable 
difference is that the illustration of the dead White- 
throated Needletail has been replaced with a represen- 


The Bird Almanac 


By David M. Bird. 2004 Key Porter Books Ltd. 70 The Espla- 
nade, Toronto, Ontario, MSE 1R2 Canada. 460 pages, Can 
$24.95 Paper. 


Within this book is a massive amount of bird-related 
information! This review could actually end right here. 
However, I will elaborate somewhat. 

Information in this book is presented in three ways: 
line drawings (restricted to the anatomy chapter), tables/ 
lists as well as glossary-style entries. These are all ap- 
propriate and lead the reader to finding information 
rather quickly. There is both birding information and 
ornithological information (and a massive amount of 
overlap which is shared by both). A glossary of over 
20 pages contains over 1000 terms — surely the word 
you're looking for must be there! 

There are simply too many categories of information 
to give more than a smattering of examples here. Both 
traditional and genetically-based classifications of bird 
families are given, followed by the massive list of all 


tation of a live, flying one. Some extra space was gen- 
erated on the main plates by moving the vagrants to a 
separate section. This is a better method of dealing 
with these rare birds. 

The introductory material has been reorganized to 
give a more logical flow. Two small sections — on DNA 
and prehistoric birds — have been replaced by more 
pertinent material. All the range maps have been up- 
dated. The result is the book is slightly smaller than 
the original, so the book remains portable in the field. 

Within the field information I noted only a few points 
I would question. The authors tend to lump species that 
more recently have been separated. These include the 
Royal and Macaroni Penguins, the Lesser and Greater 
Snow Petrels and the Yellow and Crimson Rosellas. The 
Paradise Parrot is still included, despite not having 
been seen since 1927. 

This revised edition, with the original, simpler title of 
Birds of Australia and the expansion of information, 
makes this field guide a more useful book than the 
original. 

Reference 
'Dunn, Jon L. 2002. National Geographic Field Guide to the Birds of 
North America: 4'" edition Revised and Updated. National Geo- 


graphic Society, Washington, D.C. 
Roy JOHN 


2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario 
K1J 6K5 Canada 


known bird species. Significant people are listed in sev- 
eral tables, ornithological award recipients, world-class 
listers, Taverner Cup winners, bird artists and more. 
Bird watching clubs, ornithological societies, magazines 
and journals are listed from sources around the planet. 

To compare this book with Leahy’s The Birdwatch- 
er’s Companion, a recent tome of similar intent, would 
be to have The Bird Almanac come on top. The former 
is essentially all in dictionary format, and therefore 
lacks the comparative ease (or the ease of comparisons) 
of the thematically-organized, tabular format in Alma- 
nac. There are more in-depth definitions and descrip- 
tions in Companion (it is also a much bigger book), 
and the bibliography is better- organized, but I believe 
birders will much more enjoy flipping through Almanac. 


RANDY LAUFF 


Department of Biology, St. Francis Xavier University, Anti- 
gonish, Nova Scotia B2G 2W5 Canada 


630 


2004 


Book REVIEWS 


63] 


Handbook of Birds of the World: Volume 9 Cotingas to Pipits and Wagtails 


Edited by Josep del Hoyo, Andrew Elhott and David Christie. 
2004. Lynx Edicions, Barcelona, Spain. 850 pages, illus- 
trated. Cloth $195 US. 


This volume of the Handbook of Birds of the World 
covers 819 species, including Cotingas, Manakins, 
Tyrant-flycatchers (over 50 % of the book), New Zea- 
land Wrens, Scrub-birds, Lyrebirds, Larks, Swallows 
and Martins and Pipits and Wagtails. With the publi- 
cation of this volume Handbook has now covered 60% 
of the world’s 197 families of birds. Under the current 
schedule Volume 10 will be published in 2005 and the 
final volume, 16, is due in 2011. 

It has become very difficult to review these books. 
Most reviewers, including myself, have already praised 
the content, the artwork, the scholarship, the format and 
so on. The quality of all aspects has been remarkably 
consistent. This volume is no different and it is equiv- 
alent to the other volumes. 

Volume 9 opens with an essay on ornithological 
nomenclature by Richard Banks. This is a description 
of the history, the rules of nomenclature and some of 
the pitfalls. The author has tabulated some of the most 
frequently used species names (cinereus and its vari- 
ants is the most used). He also notes that after Linnaeus, 
the originator of the nomenclature system, Philip Lutley 
Sclater, a British bio-geographer and taxonomist, has 
officially named the most birds. 

This is a timely essay for Volume 9 because it pre- 
cedes an unusual situation. Research has shown that a 
proposed sub-division of the Tyrannidae had not been 
formally recognized. Therefore the editors published a 
description of the proposed tribe as required by inter- 
national protocol. While HBOW is a little different than 
the usual scientific journal, there is no reason why it 
should not publish this text. 

The breadth of coverage starts with the wildly col- 
oured and sometimes oddball Cotingas, not just the 
dazzling Cock-of-the-Rock but the bellbirds and um- 
brellabirds too. The manakins are smaller but also bril- 
liant. [remember stopping dead at the sight of a Red- 
capped Manakin and being so transfixed I almost missed 
a much rarer flycatcher; the Sulphur-rumped. But in 
this edition the flashy birds are overshadowed by the 
“little brown jobs - LBJs.” Just under half of the 429 
Tyrannids are olive-grey-brown. Most of the larks and 
pipits are LBJs too. In fact, about 75% of the species in 
this volume fall into this category. This includes the 
dozen or so notorious Empidomax flycatchers and the 
two dozen tropical Elaenias. To separate these birds you 


need a good view plus either their song or geographical 
location to stand a chance at identification. Earlier 
this year, a colleague and I spent 15 minutes studying a 
close and very still (it was drizzling) Mountain Elaenia 
before we were satisfied with its identity. 

Even when the Tyranid is brightly coloured with a 
yellow underside and a black-and-white striped head, 
then you have 14 look-alikes to contend with. Again 
geography helps but separating Social and Lesser 
Kiskadee flycatchers can be tricky. 

One of the qualities I have learnt to appreciate more 
and more about HBOW is the perspective I get by hav- 
ing the entire world taxonomy and distribution at my 
fingertips. For example, when I encountered a new 
species, one of the eight African longlclaws (in Mac- 
ronyx) I was able to read about their relationship with 
other pipits. This despite my tendency to regard them as 
little meadowlarks (of the purely America Sturnella), 
from their appearance and habits. HBOW brings a focus 
to the biogeography and family ties to these widespread 
families. 

This access to worldwide taxonomy also raises ques- 
tions. How can the Temminck’s (Horned) Lark of north- 
ern North Africa be a different species from the Horned 
Lark of Europe and North America? The variation in 
plumage of the latter easily overlaps the paleness of the 
former (especially Eromophila alpestris bicornis, 
E.p. brandti and E.p. teleschowi). Indeed, some years 
ago, I saw a Horned Lark in Manitoba that was remark- 
ably pale and very close to Temmink’s Lark. I have 
never doubted it was nothing more than a pale version 
of our common Horned Lark. So I wonder why is Tem- 
minck’s separate from the Horned Lark, which in turn 
is split into 42 subspecies? I look forward to Volume 
10 to see how the authors handle the stonechats, in 
particular the very distinctive Madagascan Stonechat. 

The editors are continuing to maintain their uni- 
form, high standard and have settled in to a one- 
per-year production. In this volume they have made 
another useful change. Previously the plates identi- 
fied the species by a number only. You needed to turn 
to the text to match the number with a name. The 
names of all the species depicted on a plate are now 
given along with the species number in a footnote 
opposite the plate. 


Roy JOHN 


2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario 
K1J 6K5 Canada 


British Columbia: A Natural History. Revised and Updated 


Richard Cannings, and Sydney Cannings. 2004. Greystone 
Books, Douglas and McIntyre, Vancouver, British Colum- 
bia, Canada. 341 pages, 209 figures. $39.95. 


This paperback reprint is a gem. As the title indicates, 
it is a revised and updated version of a hardcover book 


first published in 1996, which won the Bill Duthie 
Bookseller’s Choice Award for the “best work pub- 
lished in British Columbia” that year, “The Lieutenant- 
Governor’s Award for Historical Writing in British 
Columbia,” and the Science in Society Award of the 


632 


Canadian Science Writers’ Association for the best gen- 
eral work. It is an informative, attractive, extremely 
well-organized book. 

The Cannings brothers (identical twins), their broth- 
er, Robert, and their late father, Steve, have each con- 
tributed masterful photographs. All four are well-known 
naturalists who have done much to encourage and fur- 
ther the study of natural history through print, radio, 
and television in western Canada. 

The Cannings’ book succeeds at a number of levels. 
Visually it is a delight, with superb, eye-catching maps 
and photographs, enhanced by informative captions. 
On initial browse, one is stimulated by the pithy, in- 
formative, boxed vignettes, such as “when is a seagull 
not a seagull?” and “Western Sandpipers.” These vig- 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


nettes will please even the most jaded of naturalists. 
Finally, the main text provides a wealth of well-organ- 
ized and thoughtfully developed information. Topics 
such as geology, the ice age, forests, mountains, grass- 
lands, and water are clearly and incisively presented, as 
the Cannings’ knowledge and enthusiasm shine through. 
British Columbia is most fortunate to have such mas- 
terful and able presenters; nature lovers everywhere 
will benefit from this book. Only superlatives apply. I 
cannot think of a single word of criticism. 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N 0J8 
Canada 


Field Guide to Bird Nests and Eggs of Alaska’s Coastal Tundra 


By T. D. Bowman, 2004. U.S. Fish and Wildlife Service 
Anchorage, Alaska. Published by Alaska Sea Grant Col- 
lege Program. 81 pages, U.S. $25.00 Paper 


This little publication is an interesting add-on for the 
field practitioner in Alaska and the arctic. This region 
includes some of the most productive bird nesting areas 
in North America, and probably in the world. It shows 
in detail nest and egg features of 70 bird species (loons, 
waterfowl and crane, jaegers, gulls and terns, owls, 
ptarmigan, alcids, shorebirds and passerines). Tavern- 
er’s Canada Goose and Cackling Canada Goose receive 
separate chapters. The species included in this guide 
were selected based on their occurrence in Alaska and 
by their “relative abundance”. Some rare species like 
Sanderling, Surfbird and Yellow-rumped Warbler are 
unfortunately not included, which will not really help 
to bring us any closer to their nest discoveries. 

This is a convenient book designed to be used in 
the field. Bird species in this book are ordered by egg 
size, which makes an interesting presentation. The book 
design and the photo arrangements are done nicely, but 
sometimes I find the egg photos a little overdone and 
repetitive; sometimes wing photos are hard to differ- 
entiate. For each species, the photos show the bird 
(often male and female), bird on the nest, the actual 
nest, eggs and (down) feathers; but this format is not 
always strictly followed. Personally, I find it somewhat 
a shortcoming that chicks are rarely presented in this 


Guide to Hawk Watching in North America 


By Donald S. Heintzelman. 2004. Globe Pequot Press, Guild- 
ford, Connecticut. 425 pages, U.S. $16.95 Paper. 


The Guide to Hawk Watching in North America, 
written by ornithologist Donald Heintzelman, is an up- 
dated version of his A Guide to Hawk Watching in 
North America, published in 1979. The new edition 
includes updated information about “hawk” (including 
vultures, eagles, harriers and falcons) biology, identi- 
fication and seasonal migration movements, as well 


guide. Of interest is the concept of a “Sizing Chart’, 
which “provides a quick reference to help reduce the 
number of prospective species to which an egg could 
belong”. Further, the book offers for geese “Parting 
Shots...” in order to identify escaping birds. A quick 
reference guide is provided for “Dark Goose Nests.” 

Six short introduction text pages provide basic back- 
ground about the species and the book concept. I ad- 
mire the challenges and amount of field work involved 
in compiling the specific nest and egg colour photos 
(over 450). A great photo collection is presented which 
includes not only, those done by the author but also 
contributions from over 70 other contributors. As I 
tend to be creative while in the field, I miss some free 
pages to write on during field work. 

The competent author has over 15 years of experi- 
ence working in the arctic, and is inspired by the 
“...dedication to improving the reliability and accu- 
racy of scientific data...”. Therefore, it can be hoped 
that these photos eventually might occur on the internet 
/www for a free use by the general and interested public. 

Overall, I think this interesting book serves it pur- 
pose well as a quick field reference and fills a vacant 
niche. I am sure it will also prove useful for Arctic 
regions of Canada as well as Russia. 

FALK HUETTMANN 

Institute of Arctic Biology, Institute of Wildlife Biology, 

University of Alaska-Fairbanks, Fairbanks Alaska 99775 
USA 


as numerous additional sites to watch raptors. The new 
edition adds a separate section on where to view Bald 
Eagles. 

The book is divided into two main parts. The first 
concentrates on general information, including species 
accounts for a wide variety of raptors other than owls, 
tips on how to identify and study hawks, a description 
of the migration seasons, an overview of different types 
of hawk watching, an introduction to field equipment 
for the activity, and an explanation of the mechanics of 


2004 


hawk flight. The second part of the book focuses on 
hawk watching sites, including U.S. hawk migration 
watch sites, Canadian hawk migration watch sites, Bald 
Eagle viewing areas, and places to view raptors out- 
side the migration season. 

The species accounts provide excellent identification 
information on new world vultures, ospreys, kites, 
hawks, eagles, harriers, caracaras and falcons. Each 
species 1s described in terms of wingspread, total length, 
field recognition features, flight style, voice, nest, eggs, 
maximum recorded longevity, food, habitat, and North 
American range. It makes for an excellent species sum- 
mary, even if it lacks references to the bird’s conser- 
vation status. 

I found the chapter on mechanics of hawk flights 
particularly interesting. Although my general under- 
standing of hawk migration was fairly good when I 
started reading this book, I had never looked into it in 
great detail. So I was fascinated to read about the influ- 
ence of general weather conditions, and the use migrat- 
ing raptors make of deflective updrafts, lee waves, ther- 
mals, thermal streets, squall lines, and leading lines. 

I was also interested to read about hawk migration 
watch sites throughout Canada, including Alberta’s 
Canmore Collegiate High School, Nova Scotia’s Brier 
Island near Digby, Ontario’s Holiday Beach Migration 
observatory near Windsor, and Quebec’s Morgan 


Book REVIEWS 


633 


Arboretum at the west end of Montreal — a sampling 
of Canadian migration watch sites with high ratings. 
Canadian entries in the Bald Eagle watch sites section 
include British Columbia’s Active Pass in the Gulf Is- 
lands, Pacific Rim National Park on Vancouver Island, 
the Fraser Valley Bald Eagle Festival near Mission, the 
Squamish Valley, the Greater Vancouver Area, and 
Prince Rupert. 

The section on other raptor viewing areas also offers 
sites in Canada, including Amherst Island and Wolfe 
Island in Eastern Ontario, both featuring Rough-Legged 
Hawks and various owls during the winter season. The 
other Canadian sites for concentrations of raptors out- 
side the migration season are the Yukon’s North Klon- 
dike Highway and South Klondike Highway. Both 
areas offer year-round opportunities to see a wide vari- 
ety of eagles, hawks, falcons and owls. 

The Guide to Hawk Watching in North America is 
an extensive and practical information source for any- 
one interested in raptor watching on this continent. 
My only objection about the book is its title, which I 
feel should be changed to Guide to Hawk Watching in 
the United States and Canada, since it lacks any ref- 
erence to sites outside those two countries. 


R. SANDER-REGIER 
RR5 Shawville, Quebec JOX 2YO Canada 


Locust: The Devastating Rise and Mysterious Disappearance of the Insect that Shaped the 


American Frontier 


Jeffrey A. Lockwood. Basic Books, New York. Hardcover. 
294 pages, Can. $39.00 Paper. 


Locust is much more than an account of a single 
species of insect. In fact, it is everything the dust cover 
promises: “A fascinating detective story” that delves 
into “history, culture, religion, and especially ecology, 
interwoven by the life story of a common insect. ...with 
vivid prose, epic thoroughness and scientific precision.” 
I would add geography and science to that list. Lock- 
wood writes engagingly. He shares his years of detec- 
tive work, providing details that only a professional 
entomologist could. His historical delvings put most 
historians to shame, and his writing skills exceed those 
of almost any living science writer. 

The book opens in Dodge County, Nebraska, in July 
1875, a drought year, with clouds of locusts obscuring 
the sky, their wings crackling like a horrific blaze. 
Limbs of willows bent to the ground under the weight 
of the insects, as adjacent cornstalks were stripped 
bare. When a sheet of insects six inches thick passed 
over a perpendicular ledge of rock, they caused a roar- 
ing noise similar to a cataract of water. In Utah, locust 
eggs were counted at 743 million eggs per acre. The 
voracious insects literally ate the clothing off human 
limbs, then entered homes to eat objects such as win- 
dow blinds. When domestic chickens gorged on the 
locusts, their eggs and flesh became inedible. Farmers 
and their families lost their gardens and crops, and 


were on the brink of starvation. 

The U.S. Army under General Ord saved many farm 
family lives by far exceeding normal army routine. 
Ord issued thousands of infantry coats, shoes and mil- 
itary blankets, as well as large amounts of army rations. 
Lawmakers apportioned money to distribute wheat seed 
for planting the following spring. Without Ord, thou- 
sands would have died. 

North America was blessed with hundreds of species 
of grasshoppers, but only a single species of locust, the 
Rocky Mountain locust, Melanoplus spretus, named 
by Benjamin Dann Walsh back in 1866, caused such 
vast destruction. Ingenious but rather ineffective ma- 
chines such as suction machines and flame throwers 
were invented to combat the locust; two men and a team 
of horses could incinerate ten acres of locust-infested 
fields in a day. One Minnesota community alone had 
a thousand coal-tar hopperdozers that could harvest 
150 000 locusts per hour. 

On the scientific front, three entomologists did their 
best to help combat the locust. Charles Valentine Riley 
was the state entomologist for Missouri. Cyrus Thomas 
and Alphaeus Spring Packard, Jr., held similar posts 
in Illinois and Massachusetts. Riley was able to show 
that the locust had the potential to increase its popu- 
lation 100-fold from one generation to the next; he 
argued that for every bushel of locust eggs destroyed, 
100 acres of crop could be saved. He founded the Na- 


634 


tional Insect Collection and through his efforts a branch 
unit in economic ornithology arose. This office became 
the Bureau of Biological Survey and later metamor- 
phosed into today’s U.S. Fish and Wildlife Service. 

Locust has important Canadian content. Entomolo- 
gist Norman Criddle of Manitoba devised a widely- 
used mixture of copper acetoarsenite, molasses and 
horse manure to combat later outbreaks of somewhat 
less harmful grasshoppers. Criddle also collected the 
world’s last two specimens of Melanoplus spretus on 
19 July 1902. Three Saskatchewan men, Paul Riegert, 
Bill Chapco, and Bob Randell, also helped solve prob- 
lems discussed in later chapters. 

Why did the Rocky Mountain grasshopper become 
extinct? Five different theories in turn held sway: the 
spread of alfalfa; the demise of the Bison; changes in 
weather; overgrazing of grasslands; widespread prairie 
fires — but each was discredited, though two were later 
revisited as contributing factors.. 

To allow DNA studies and do radiocarbon dating, 
Lockwood determined to find locust specimens en- 
tombed in glacial ice for a century or more. His first 
try, in 1987, yielded a quarter pound of dried grass- 
hopper parts; when results were submitted to a leading 
entomological journal, the editor rejected the paper 
and informed Lockwood that “you have mistaken 
natural history for science.” In 1988, Lockwood and 
colleagues collected 134 specimens of twenty species 
of grasshopper, but no locust, on the glacier. In 1989, 
they obtained 4 mg of grasshopper parts, including 
mandibles that appeared to match those of extant pre- 
served specimens of Melanoplus spretus. Finally, in 


Mammals of the World: A Checklist 


By A. Duff and A. Lawson. 2004. Yale University Press, P.O. 
Box 209040 New Haven, Connecticut 06520-9040 USA. 
312 pages, U.S.$45.00. 


Perhaps I should have bought this book some time 
ago, or at least its predecessor; Mammal Species of 
the World (Edited by D. E. Wilson, and D. M. Reeder, 
1993, Smithsonian Institution Press), which contains 
the names of the recognized species of mammals doc- 
umented at that date. So many times I have struggled 
to understand which species are present in a specific 
area. With birds this is fairly easy to resolve. English 
names are reasonably standard and scientific names 
are the ultimate guide. With birds I have rarely had to 
probe recent literature to catch up on the latest splits. 
With mammals this is much more difficult. It took 
some time to clarify the species and distribution of 
fur seals. The confusion of English names for wilde- 
beest left me very perplexed. My latest book on African 
mammals was particularly mystifying. It is a transla- 
tion from German to English and uses different Eng- 
lish names for the plates and the text. The scientific 
names are not current. Starting with this book it took 
a lot of work to create a list of lemurs that showed the 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


1990, at the melting edge of glaciers, they collected 250 
locust bodies, including 14 males with well-preserved 
abdomens and genitalia, allowing unequivocal identi- 
fication and DNA analysis. These insect bodies had 
taken about 150 years to travel 300 m as the ice moved 
from the crevasses where they had been entombed. 

Why, indeed, had Melanoplus spretus become 
extinct? For once, the unplanned effects of human activ- 
ity had an effect for the better. Lockwood explains that 
the “base locality” of the locust was, between out- 
breaks, restricted to a few relatively small areas in val- 
leys within the Rocky mountains, where eggs could 
be deposited in sand and gravel. Following European 
settlement, floods occurred more often due to overgraz- 
ing on the slopes, irrigation periodically flooded the 
valleys, and the locust eggs lost their vitality. Plough- 
ing and harrowing destroyed the eggs. The new alfal- 
fa crops were inimical to development of the locust 
nymphs. The enigma has been solved. 

Lockwood is a consummate writer. His eminently 
readable book is a detective story, keeping the reader in 
suspense to the final chapter. Admittedly this review 
has given away the ending and thus has spoiled some 
of that suspense, but the pleasure of this book lies more 
in its details and insights than it does in the suspense. 
I strongly recommend this book to everyone with an 
interest in a good story well told. 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N 0J8 
Canada 


currently known species and sub-species. With Mam- 
mals of the World this exercise is simple. 

This current list contains 5069 species. From the 
Wilson and Reader list of 4629 the authors have sub- 
tracted 41 species that were extinct prior to 1800. These 
include such creatures as Steller’s Sea Cow, hunted to 
extinction by 1768. More recently extinct species are 
left in, presumably as, in theory, there is a chance of 
still finding one alive. The Thylacine comes to mind. 
Duff and Lawson have added 522 “new” species. A 
large portion of these come from two sources. Juliette 
Clutton-Brock’s work on domestic animals has elevat- 
ed numerous creatures to species status (the Domestic 
Pig goes from Sus scrofa to Sus domesticus for exam- 
ple). The second source of change is the assignment of 
species status to numerous sub-species (Colin Groves 
of the Australian National University has been active in 
this exercise). Finally, 41 species have been demoted in 
synonymy. Sadly this loses us the Queen of Sheba’s 
Gazelle (now a subspecies of the Arabian Gazelle), a 
romantic loss at least. The changes in all the above 
categories are fully explained in the appendices. Also 
included are the nomenclatural changes in scientific 
names. 


2004 


The authors say they have followed Wilson and 
Coles (Common Names of Mammals of the World. 
2000. Smithsonian Press) version of the English names 
with corrections and conversion to “well-established 
names.” Generally they do not list alternative names 
so an animal like the Cougar, or Mountain Lion, or 
Catamount only gets listed as Puma. When I used the 
index, being uncertain of the author’s choice, I looked 
up Lion (Panthera leo) knowing that Cougar would 
be nearby. The most odd name I found was Sewellel, 
the Chinook Indian name for the Mountain Beaver. 
This was the first time I had seen this name although 
Audubon used it on his painting of Aplodontia rufa. 

Naturally I compared the list of lemurs that I had 
created to that of Duff and Lawson. I was not surprised 
to find many that I had as sub-species had been elevat- 
ed to full species. Otherwise the lists were the same 
except for Grey-brown Mouse Lemur (Microcebus 
griseorufus). R. M. Rasoloarison, S. M. Goodman, and 
J. U. Ganzhorn first described this cute, hamster-like 
lemur in 2000. I have since been fortunate to see this 
little beast in South-western Madagascar. This omis- 


BooK REVIEWS 


635 


sion is surprising as the author’s references go up to 
2002, but to be fair this is a family undergoing con- 
stant change. 

I was also surprised that Canada was not mentioned 
in the range of the Red Wolf (Canis rufus). John The- 
berge’s work has shown that Algonquin wolves are 
closer to Red Wolves than the more common Timber 
Wolf (Canis lupus). This information is far more wide- 
ly known. 

For those of us that like to travel and see mammals 
in their native habitat this book is a good and useful 
guide. However, it is only a list and, although it will 
not resolve all taxonomic questions, it will bring 
clarity and order to your research and therefore it is a 
valuable reference book. As it is based on Wilson and 
Reeder’s out-of-date book it is more up-to-date, but I 
understand a revised edition of Wilson and Reeder 
will be published soon — should we wait? 


Roy JOHN 


2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario 
K1J 6K5 Canada 


Parental Behavior in Lepidosaurian and Testudinian Reptiles: A Literature Survey 


Louis A. Somma, 2003. Krieger Publishing Company, Mal- 
abar, Florida. x + 184 pages, U.S.$33.50 Cloth. 


This book is exactly what it purports to be. It is a 
summary and exhaustive bibliography of literature on 
parental behaviour in lizards, snakes, amphisbaenians, 
tuatara and turtles. Passing reference is also made to 
literature on other vertebrates including dinosaurs. 
This book focuses exclusively on behaviour as opposed 
to physiological adaptations and so, while parental 
behaviour in some species of snakes and lizards is well 
known, many readers will be astonished to find turtles 
included. Yet Somma cites references that provide 
some evidence of parental behaviour for seven species 
of turtles. 

Fifteen categories of parental behaviour are discussed 
including defence, thermo- and hydro-regulation, assis- 
tance during hatching and facilitated feeding. For those 
unfamiliar with research in this fascinating field the 
range and diversity of parental behaviours in these 
reptiles will intrigue and astonish you. Unfortunately, 
description and discussion of these behaviours is lim- 
ited to the first 11 pages of the book. This is followed by 
46 pages of tables summarizing the results and guiding 
the reader to literature on various topics and taxa. The 
remainder of the book consists of 100 pages of refer- 


ences and an index to subjects and taxa. Readers should 
be aware that the index is inconsistent as to whether 
it includes references to taxa within the tables (for 
example, it does for Trachemys stejnegeri malonei but 
not for Eumeces fasciatus). 

The primary purpose of the book seems to be to raise 
the profile of parental behaviour in non-avian reptiles 
and encourage further research on this understudied 
phenomenon. It is unfortunate that having read such an 
enormous amount of information on this topic, Somma 
does not provide more insights into its evolution and 
ecological ramifications but perhaps the field is too 
young for such a synthesis. Certainly for anyone inter- 
ested in embarking on research into this topic it is an 
invaluable and relatively inexpensive reference. For 
those who are most keen on this topic, Somma has also 
published an addendum to the book (Somma, 2003). 


Reference 

Somma, L. A. 2003. Parental Behaviour in Lepidosaurs and Turtles: 
Source Addendum. Bulletin of the Chicago Herpetological Soci- 
ety 38(4): 65-76. 


CAROLYN SEBURN 


7210 Clarenda Street, Ottawa, Ontario K2B 7S5 Canada 


Prairie Ghost: Pronghorn and Human Interaction in Early America 


By Richard E. McCabe, Bart W. O’Gara, and Henry M. 
Reeves. 2004. 176 pages, U.S.$29.95 Cloth. 


The Pronghorn, Antilocapra americana, formerly 
called “antelope,” is the “most American” of the con- 
tinent’s terrestrial wildlife, since it is found nowhere 
else. It is the world’s second fastest land animal, and 


perhaps the most inquisitive. Protuberant eyes allow it 
a nearly 360-degree field of vision. Large lungs, heart 
and trachea permit it to achieve great speed. 

This historical look at the Pronghorn is thoroughly 
researched, with informative tables, extensive refer- 
ences, and well-chosen, sumptuous illustrations. It 


636 


delves deeply into anthropology. The Pronghorn rep- 
resented fleetness, alertness, and pertinacity, and was 
a catalyst in the social structure and welfare of most 
plains tribes. It is no wonder that many place names 
today derive from this one species, but who would have 
expected 108 such instances in Arizona and 79 in Wy- 
oming? Two detailed appendices list dates of eyewit- 
ness accounts between 1540 and 1896, and the names 
of the Pronghorn in each native language. Historical 
accounts go back to Sahagun in 1569 and Hernandez 
in 1651, both in Mexico. Lewis and Clark popularized 
it during their exploratory journey of 1804-1806, and 
George Ord gave the Pronghorn its binomial Latin 
name in 1815. 

Before Europeans reached the Americas, aborigines 
hunted the swift-footed Pronghorn with three instru- 
ments: sling, atlatl, and bow-and-arrow. A well-condi- 
tioned, determined native could sometimes, in spite of 
his much slower pace, outlast and eventually tire and 
kill a Pronghorn. More often, stalking, pursuing, sur- 
rounding, luring, calling, impaling, ambushing, netting, 
driving with v-shaped fences, setting prairie fires, and 
making pitfalls, were the methods used to kill them 
for food. Pronghorn hunting required more prepara- 
tion and more co-operative effort than did hunting of 
the larger Bison. The Pronghorn was then one of the 
natives’ most important food sources, especially where 
Bison were scarce near the edge of their range. An 
antelope skin was thinner and lighter than that of a 
Bison, and thus more suitable for clothing. A native 
would obtain about 45000 calories from a 43-pound 
Pronghorn carcass. 

Pronghorn bones were used as toys, rattles, awls, 
pipes, fishhooks and decorations. Sinew served as 
strings for bows, and for sewing. A skin stretched tight- 
ly over a section of hollowed tree formed a drum. 
Marrow was rubbed on sunburned or chapped lips and 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


skin. A Pronghorn fawn was used as bait on branches 
above a pit where an Indian patiently waited to catch 
a Golden Eagle for its feathers. Images of the Prong- 
horn were used to decorate native pottery, on the walls 
of caves at ten known sites, and as effigies. 

Hunting became much easier after guns spread north 
from Mexico and guns and ammunition were obtained 
in trade, but this soon led to squandering of the once- 
precious resource by a veritable army of 5000 white 
hunters. From 1874 through 1877, more than 100 000 
Pronghorn hides were shipped from the plains annually. 
As the Bison almost vanished, the Pronghorn became 
even more important as a food source, and their num- 
bers dwindled as well, from about forty million to fewer 
than 15000 in 1910. Bereft of their two largest natural 
food sources, native peoples were overwhelmed, sub- 
jugated, displaced, and pauperized of their culture, iden- 
tity and social options, losing much of their spirit and 
vitality. 

Although three of the four last sections tell a grim 
story, dealing with the near demise, in turn, of the 
Bison, the people, and the Pronghorn, the book ends on 
a positive note with the birth of the conservation ethic. 
In 1887, Theodore Roosevelt convened a dinner meet- 
ing in Manhattan to launch the Boone and Crockett 
Club; he and the other founders had a strong affection 
for the Pronghorn, which eventually benefitted from 
the conservation ethic and philosophy of this fledgling 
movement. 

This attractive, scholarly, modestly-priced book 
belongs in major libraries. It would make a perfect gift 
for anyone interested in history, geography, anthro- 
pology, or big game hunting on the plains. 


C. STUART HOUSTON 


863 University Drive, Saskatoon, Saskatchewan S7N 0J8 
Canada 


A Guide to the Birds of St. Helena and Ascension Island 


By N. McCulloch. 2004. The Royal Society for the Protection 
of Birds, The Lodge, Sandy, Bedfordshire, SG19 2DL 
Great Britain. iv + 92 pages, 11 GBP. 


The booklet is intended for the visitor and has three 
parts. The first is an overview of island history from 
time immemorial to present. The second is a site guide 
giving where to find birds on these islands. The last 
part is the bird accounts. 

The bird accounts cover 31 species for St. Helena 
and 46 for Ascension that a visitor is likely to see in a 
day trip around each island. Of these 11 are seabirds 
and 10 are shorebirds on St. Helena and 13 are seabirds 
and 16 shorebirds on Ascension So only about 30 % of 
the island species are land birds, with most of them 
being introduced. There have been 41 other attempts 
at introduction on St. Helena and 9 on Ascension 

The account of the history is fascinating and so 
typical of remote islands, especially after its “discov- 
ery” by humans. This is not good bedtime reading 


though. The destruction of the islands ecosystem is the 
usual unpleasant tale of wanton destruction misguided 
mistake and foolish carelessness. There have been 41 
failed attempts to introduce everything from a white-eye 
to an ostrich to St. Helena and 9 attempts on Ascension. 
This includes the introduction of House Sparrows in 
1986 (to be company for a ship-assisted vagrant who 
arrived in 1985). Will we never learn? On a happier note 
there are programs underway to restore habitat for 
around 40 species of endemic plant and 9 species of 
creepy crawlies (one — the Golden Sail Spider — is 
illustrated) as well as birds. There are also programs 
to reduce cats, goats and the like. 

The bird finding section is typical of current field 
guides, portraying the bird on the left and giving infor- 
mation on the right. It covers 28 species. Half of these 
are seabirds. The remainder are land birds, almost all 
introduced. The English names are fairly consistent 
with other texts, but there are no references to alter- 


2004 


| native names. For example Maderan vs. Band-rumped 
Petrel or Parasitic Jaeger vs. Arctic Skua. The text here 
along with the rest of the book is good and clear. 

The author has added a table of accidental species 
for the two islands (31 on St. Helena and 46 on Ascen- 
sion. This did not quite match my own list, but I think 
the difference is due to the political boundary that in- 
cludes Tristan de Cunha and Gough Islands, thus add- 
ing several Antarctic species) 

The artwork consists of watercolours that are refined 
sketches of birds during their normal activities. This 
gives a better idea of their jizz than the more formal- 

ized field guide style of the recent books, and is more 
_ akin to the style of postwar books. Indeed, my favour- 
ite is a perky field sketch of a Java Sparrow that fronts 
| the section of land birds on which you can almost 
| count the small number of brush strokes. The reader 
| can compare this with the more “finished” plate in the 
accounts section. 

I found it ironic that the endemic Madagascar Fody 
| was introduced, not from Madagascar but from Mau- 
| ritius (where it is an introduced threat to the endemic 


| BOTANY 


BooK REVIEWS 


637 


Mauritius Fody) and became so numerous it was part 
of the St. Helena cage bird trade. How convoluted we 
make the world. 

So why go to these remote places? First there is the 
attraction of wild oceanic islands. While they are no 
longer the lush paradise first seen by the Portuguese 
they still are dramatic. The rich brown cliffs surround- 
ed by blue sea set off the white, guano-capped islands. 
Second, however diminished, there are good seabird 
colonies; always exciting places. And last, for the hard 
core, they are the only places to see Ascension Island 
Frigate and Wirebird. While remote, these islands are 
not inaccessible. There are some tours that include them 
on a cruise, although these are very expensive. Routine 
commercial sailings leave from Cardiff, Wales and Cape 
Town, South Africa a few times a year. They dock in 
St. Helena and Ascension for a day or two — enough 
time with this guide to see most of the islands birds 


Roy JOHN 


2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario 
K1J 6K5 Canada 


| Flora and Climatic Conditions of the North Pacific: A Collection of Scientific Papers 


| Edited by A. N. Berkutenko, H. G. Lumsden, and D. Lumsden. 
2001. Institute of Biological Problems of the North, North- 
East Scientific Center, Far East Branch, Russian Academy 
of Sciences. Magadan. 189 pages, No price available. 


The subtitle — A collection of papers — better indi- 
| cates the contents of this book than does its lead title. 
| What we have here is a nicely produced small book 
| consisting of papers on flora and vegetation, for the 
| most part, but also one paper each on fungi and eth- 
nobotany, two on seed biology, and one on the influ- 
| ence of air masses from the Sea of Okhotsk on sum- 
mer temperatures in Japan. An odd mix, but perhaps 
| the inevitable result of the need to collect sufficient 
| papers, achieve critical mass, and gain publication. 
| To one interested in floristics and taxonomy, several 
_of these papers are informative and useful. Since the 
| book is entirely in English, it opens to a wider audi- 
ence than usual the results of botanical studies in the 
Russian Far East. The prime mover for this collection 
was A. N. Berkutenko who wrote entirely or con- 
_ tributed to six of the collected 15 papers. Since it is 
difficult to generalize the disparate contributions, I 
will give a précis of each. 
Yakuboy et al. provide a brief sketch of the physical 
_ setting, a history of botanizing, and an annotated check- 
| list of 235 species for the flora of Avachinsky volcano. 
| The next paper by Mochalova describes the very 
small flora of very small islands of the Commander 
archipelago and the effects of bird colonies on the veg- 
etation there. Two papers by Khoreva and by Berku- 
tenko et al. discuss the vegetation and flora of two 
islands in the Sea of Okhotsk and provide checklists 


for the Yams Islands and Nedorasumenia Island, the 
latter with the unexpected occurrence of the Asiatic 
shrub Caragana jubata. 

Sinelnikova gives a synopsis of the plant cover and 
a checklist for 454 species of vascular plants found at 
the Orotuk field station in the upper Kolyma River 
region. The station lies in larch taiga near the Koly- 
ma floodplain. The Chosenia arbutifolia and Populus 
suaveolens of the gallery forest and the Pinus pumila 
communities farther upslope are among the memorable 
botanical images I took away from my trip to the 
region. A short paper by Berkutenko and Khoreva, in 
a structure now familiar, provides a sketch of the veg- 
etation and a checklist to the 98 species found at the 
Mount Kamenny Venets nature monument, which in- 
cludes the endemic willow Salix magadanensis. Thus 
ends the first 117 pages. From here on, the papers have 
less to do with each other or to what has gone before. 

A list of 161 macromycetes in the Magadan Preserve 
by Sazanova is followed by a report by Berkutenko 
and Yukawa of the first record for the orchid Liparis 
kumokiri for mainland Russian Far East.Next Misako 
proposes that morphological variation in the Japanese 
Sanguisorba tenuifolia has originated from hybrids 
between S. parviflora and S. officinalis occurring in 
coastal Russian Far East. In a brief, idiosyncratic but 
interesting essay, Berkutenko contrasts with ethnobotan- 
ical anecdotes the changes that have taken place in 
people’s live since Krasheninnikov reported on 18" 
century habits of the local people on Kamchatka. She 
includes some differences between practices in the 
Russian Far East and Alaska as well. This paper is fol- 


638 


lowed by comments on the germination of 78 species 
by Andrianova and Berkutenko: which taxa require 
pretreatments and which kinds of treatments are effi- 
cacious. A paper by Kryukov reports briefly on the 
relative germination success of 27 species following 
different periods of storage. Haese describes a phytoso- 
ciological analysis of coastal tundra using the methods 
of Braun-Blanquet, and provides tables typical of this 
analysis. Hanno and Oka in a short paper that reads 
more like a proposal noted that the Sea of Okhotsk 
affects the climate of eastern Japan. Pachomov and 


ENVIRONMENT 


The Russian Far East 
By Josh Newell. Second Edition, 2004. Daniel and Daniel, 


Publishers, McKinleyville, California, USA. 486 pages, 
U.S.$59.95. Paper. 


This is a massive and thorough compendium. The 
subtitle reads, “A reference guide for conservation and 
development.” The format is a series of chapters each 
concerning one of the major administrative divisions 
of the huge region, abbreviated as “RFE.” These divi- 
sions are: Primorsky Krai, Khabarovsk Krai, Jewish 
Autonomous Oblast, Amur Oblast, Republic of Sakha, 
Magadan Oblast, Chukotsky Autonomous Okrug (Chut- 
kotka), Koryak Autonomous Okrug (Koryakia), Kam- 
chatka Oblast, Sakhalin Oblast. There are over 50 maps, 
and a host of tables, figures and photographs, as well 
as an index. 

Vegetation is organized into the classic Tundra and 
Taiga formations. This book divides Tundra into two 
parts: “Arctic Tundra” and “Tundra” (referred to in 
much Russian ecological literature as “High Arctic” 
and “Low Arctic.”) Taiga is defined correctly as “the 
large mass of the boreal forest that forms the heart of 
the RFE.” The southern taiga, which in much of Russ- 
ian ecological literature is known as Ussuri Taiga, is 
also known as “Dark Taiga” because of the high per- 
centage of spruce and pine. 

This division serves to differentiate it from the 
northern part of the Taiga which is widely known in 
the Reindeer literature as “Light Taiga’” because of the 
very large percentage of the forest cover of deciduous 
larch. The Reindeer literature, and much ecological 
literature, also designates the southern-most tundra, 
combined with the northern-most taiga as ‘‘Forest- 
Tundra.” This recognition of it as a separate entity is 
undoubtedly because the Forest-Tundra is particularly 
important as winter pasture for the semi-domesticated 
Reindeer. 

The Ussuri taiga is relatively familiar to English- 
speaking biologists because of translations of works by 
such authors as Sdobnikov and Arseniev, as well as 
Kurosawa’s famous 1975 film about Arseniev and 
Dersu. The number and distinctiveness of the species 
of plants and animals of the RFE is legendary, espe- 
cially the Ussuri taiga. 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Sinelnikova present six years of observations on the 
effect of artificial warming on the growth of four tun- 
dra plants at or near the field station described by 
Sinelnikova (see above). I fear these studies in support 
of the International Tundra Experiment (ITEX) proj- 
ect will be lost to the [TEX community in these pages; 
it belongs with others of its ilk. 


DAvID F. MURRAY 
University of Alaska Museum of the North, Fairbanks, Alaska 


Each of these 10 administrative regions receives a 
section of the book. Each begins with an overview, 
ranging from a paragraph or so to a number of pages, 
followed by extensive entries on: location, size, climate, 
geography and ecology (including carbon stocks), major 
ecosystems, protected areas and their problems, bio- 
diversity hotspots, political status, natural resources, 
main industries, infrastructure, foreign trade, economic 
importance in the Russian Federation, general outlook. 

This latter section is particularly interesting for such 
items as a full-page table of major environmental issues 
and problem areas of each of the regions: fishing, 
energy, timber, mining, agriculture. There are pertinent 
discussions of the weaknesses of various Russian gov- 
ernmental regulations concerning oil, gas and mining 
operations compared to the (already-weak) United 
States and Canadian regulations. 

There is discussion of various schemes with poten- 
tially-severe effects as well as prospects for a sustain- 
able economy, ENGOs, the UN Global Environment 
Facility, foreign government aid agencies, other prom- 
ising sectors such as ecotourism and NTFP (Non Tim- 
ber Forest Products). There is rather detailed consid- 
eration of the degradation of zapovedniks (“protected 
areas”) by tourism, poaching (salmon, caviar), bear 
killing (for paws and gall bladders). 

The forests of Kamchatka are particularly vulnerable 
and critical for mitigating floods and protection of 
salmon spawning grounds. (Remember British Colum- 
bia?). In the RFE the largest emitter of atmospheric 
CO, is fossil fuels combustion, but second place is de- 
forestation leading to loss of carbon-rich boreal forest 
and replacement by pioneer types of forests and shrub- 
by vegetation. More than 90 percent of the logging in 
the RFE is by clearcut. Even plantation-forests do not 
recover pre-logging stocks of CO,; managed planta- 
tion-forests usually contain only 4 to 4 the carbon 
that undisturbed forests do. 

Activities to “Save Tropical Forests” are having 
adverse effects on the taiga of the RFE... “Plywood 
manufacturers are promoting Russian larch (tamarack) 
as a green alternative to tropical luan timber and have 
been steadily increasing levels over the past decade... 


|More than 98 percent of all Russian larch grows on 
/some form of permafrost (continuous, discontinuous, 
| sporadic) making large-scale logging of the species an 
| unwise proposition” (page 31). 

“The sheer size of the Siberian and RFE forests and 
| the diversity of their plant and animal life and habitats 
| make these forests a tremendously important factor in 
|}Russia and the world. Loss of habitat, mostly from 
| forest exploitation, fire, disease and inappropriate man- 
| agement is the most serious threat to the unique bio- 
diversity of climax forests. The survival of endangered 
| species, such as the Siberian Tiger ... depends on the 
/maintenance of large, undisturbed forest areas. Frag- 
‘ile permafrost areas must be recognized as environ- 
‘mentally critical and the forests’ large contributions to 
carbon sequestration must not be jeopardized. Such 
environmental considerations are currently inadequate- 
ly incorporated in the planning process or not properly 
) addressed in forest management and harvesting activ- 
|ities. As a general requirement for attending to these 
environmental concerns large forested areas must re- 
‘main undisturbed and forests outside protected areas 
/need appropriate and complementary management.” 
(page 32) 

The Russian Far East has the endangered Siberian 
Tiger (Panthera tigris altaica) while Canada has the 
}endangered Woodland Caribou (Rangifer tarandus 
caribou) (Pruitt and Baskin 2004; Schaefer 2003). Pages 
35 — 39 also bring problems in the RFE directly home 
to Canada: “A major problem is the lack and loss of 
| field scientists and field workers because of dramatic 
cuts in government funding...” We can compare the 
situation in the RFE with the massive cuts in scientif- 
ic staff of Canadian Wildlife Service, National Parks 
| Canada, Canadian Meteorologial Service and the Cana- 
| dian Museum of Nature, beginning with the Mulroney 
Conservatives and not restored by later governments. 
In Canada, the problem is compounded by the growing 
infatuation of university biology, botany and zoology 
| departments with “computer models” or “keyboard 
ecology” instead of learning about real animals and 
| plants (Ehrenfeld 1993; Futuyma 1998; Noss 1998). 

__ The book is almost overwhelming in its information: 
history of European invasions, lists (with comments) 
of major species of plants and animals, lists of endan- 
gered species (usually with comments regarding threats 
to survival), lists of protected areas by type, size, date 
of establishment, biodiversity hotspots, human econ- 
-omy and environmental impact. 

The sections on Indigenous Peoples and their decline 
_in numbers, relations and their problems with “modern” 


BooK REVIEWS 


639 


extractive economy (mining, logging, etc.) is particu- 
larly instructive because of the great resemblance to 
the history of similar Aboriginal groups and their ex- 
ploitation or rejection by Canadian governments and 
industries. 

We read much in the western press about the wide- 
spread pollution and environmental destruction dur- 
ing the Soviet era, but “While the focus has been on 
nuclear, air, and water pollution and, somewhat less, on 
the wanton waste of resources caused by inefficient 
production, scholars have largely ignored what may be 
the most significant environmental legacy from the 
Soviet era, and what is Russia’s greatest legacy to the 
planet: wilderness.” (page 29). 

Although the Russian Far East is well over the shoul- 
der of the world from Canada, the unity of the taiga 
(so-called “boreal forest’) is evident by the similarity 
of its animals and plants to those in Canada. We can 
also see in this compendium disturbing similarities to 
the problems affecting the Canadian taiga (Pruitt and 
Baskin 2004; Schaefer 2003). Comparison of amelio- 
rating efforts will be valuable. There are many places 
in Canada which should have copies of this valuable 
compendium: Federal and Provincial departments of 
Conservation, Natural Resources, Forestry, Wildlife, 
Aquatic Resources, Fisheries. All University libraries, 
the libraries of all ENGOs concerned with conservation 
and sustainable use of renewable resources, anthropol- 
ogy and aboriginal activists, everyone teaching Field 
Biology, Ecology, Boreal Ecology, Resource Manage- 
ment, Forestry, Wildlife Management, Fisheries Man- 
agement or Sustainable Development course or seminar 
should have access to a copy. 


Literature Cited: 

Ehrenfeld, D. 1993. Beginning again (pages 65-72). Oxford, Oxford 
University Press. Pages xiv + 216. 

Futuyma, D. 1998. Wherefore and whither the naturalist? American 
Naturalist 151(1): 1-6. 

Noss, R. F. 1998. Does conservation biology need natural history? 
Wild Earth 8: 10-14. 

Pruitt, W. O., Jr., and L. M. Baskin 2004. Boreal Forest of Canada and 
Russia/Taiga of Canada and Russia. Bilingual, English and Russian, 
parallel texts. Pensoft Publishers, Sofia, Bulgaria, illustrated: 163 
pages ISBN 954-642-199-5. 

Schaefer, J. 2003. Long-term range recession and the persistence of 
Caribou in the taiga. Conservation Biology 17(5): 1435-1439. 


WILLIAM O. PRUITT, JR. 


Department of Zoology and Taiga Biological Station, Univer- 
sity of Manitoba, Winnipeg, Manitoba R3T 2N2 Canada 


640 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Resource and Environmental Management in Canada: Addressing Conflict and Uncertainty 


Edited by Bruce Mitchell. 2004. Third Edition. Oxford Uni- 
versity Press, Don Mills, Ontario, 608 pages. Can $46.50 
Paper. 


Canada is a resource-rich country. The nation’s his- 
tory and folklore are a story of resource exploitation 
and the people who trapped, fished, farmed, mined and 
lumbered. In many ways, much has not changed as 
evidenced by the national wealth accrued from natural 
resources, the dependence of many towns and regions 
on primary resource use, and the growing social con- 
flicts centred on the type and intensity of resource use 
and conservation. 

It is generally acknowledged that all natural ecosys- 
tems are more or less human-dominated. Recent, well- 
cited studies have estimated that humans appropriate 
over 30% of the total terrestrial net primary produc- 
tion. This is a remarkable level of consumption for 
humans that represent roughly 0.5% of the total het- 
erotroph biomass on Earth. Spatially-explicit studies 
have identified appropriation levels that range from 
> 6% for South America to > 80% for south-central 
Asia. North America falls in between with a human 
appropriation level of 23%. 

Given these levels of human domination of the 
earth’s resources, it is understandable that the notions of 
conflict and uncertainty would be chosen as thematic 
foci for a book on resource and environmental man- 
agement; conflict among differing values and ideolo- 
gies, and uncertainties over adequate scientific and 
social knowledge. 

A long-standing textbook, now in its third edition, 
Resource and Environmental Management is intended 
primarily for undergraduate university students. The 
book’s 20 chapters are broken down into three parts: 
(1) emerging concerns, (2) enduring concerns, and (3) 
contemporary responses to these concerns. Emerging 
concerns include ecosystem health and ecological 
integrity, globalization and neo-conservatism, First 
Nations and resources, feminist perspectives, climate 
change, and water security. Enduring concerns focus 
on traditional resource issues such as fisheries, agricul- 
ture, forestry, wildlife management, mining, and parks 
and protected areas. The third part addresses some of 
the approaches used to develop resource and environ- 
mental visions. Among others, these include search 
conferences, ecosystem approaches, adaptive manage- 
ment, environment impact assessment, participatory 
approaches, governance issues, and environmental jus- 
tice. The majority of contributing authors are geogra- 


phy and environmental studies academics. 

In any collection of contributed essays, consistency 
in style and quality is always a challenge. Conflict and! 
uncertainty act as foundational themes and tie together 
all chapters. Most, if not all authors, attend to this 
theme and regularly make reference to other chapters 
in the book. Many of the chapters have been rewritten 
and updated for this edition and seven new chapters; 
have been added. The editor has prefaced each major; 
section with a good synthesis and summary of each) 
chapter thus helping the reader to situate the individ- 
ual contributions. 

Authors provide a sound scientific and technical 
analysis of each resource issue, generally up-to-date 
resource statistics, and policy implications. The strength 
of the edited volume is the authors’ attention to the 
social, cultural and political dimensions of resource 
and environmental management. A former forest ecolo- 
gy professor of mine was wont to proclaim: “Forestry 
is more about people than it is about trees.” The same 
can be said for all other resource issues. Because of the 
essential human dimension, ecosystem-based natural 
resource management is not rocket science — it’s much | 
more complicated than that. The amalgam of biophys- 
ical, cultural and social complexities fuel conflict and 
promote uncertainty. This inherent complexity of re- 
source and environmental management is highlighted 
throughout the text. Well discussed, as well, are a 
number of significant mechanisms that attempt to deal 
with the often intractable and inherently stochastic 
dimensions of institutional human behavior. 

I found some of the essays unnecessarily long- 
winded and somewhat pedantic. Twenty chapters weigh- 
ing in at over 600 pages do not make for easy or nec- 
essarily enjoyable reading, especially given the diversity 
of chapter topics. The chapter on forest management 
is noticeably remiss in addressing contemporary sus- 
tainable forest management paradigms. No mention 
is given, for example, of the current developments in 
the emulation of natural forest landscape disturbances, 
a key concept in sustainable forest management. As 
an edited book it does not possess the thrill or the 
fast-paced narrative of a single author. As a textbook, 
it has all the information, but you may find yourself 
simply dipping into chapters of specific interest. 


JOHN MCCARTHY 


Holy Rosary Parish, 
NIE 1V6 Canada 


175 Emma Street, Guelph, Ontario 


d 


2004 


| MISCELLANEOUS 


By Jonathan A. Edlow. Second Edition, 2004. Yale Universi- 
ty Press, New Haven and London. 304 pages. U.S.$29.95 
Paper. 


There have been few newly-emerging infectious dis- 
;eases that have generated as much controversy as to 
their etiology and treatment as Lyme disease. From 
the “conventional” viewpoint, the disease is easily and 
accurately diagnosed, and short-term antibiotic treat- 
ment is effective; in the “alternative” point of view, 
‘the diagnosis of late-presenting cases is often missed 
_ due to inaccurate diagnostic tests, and long-term antibi- 
_ Otic treatment of such cases is necessary for a cure. This 

readable book gives a fair and balanced account of the 
differing points of view of these two camps. It covers 
| the history of the discovery of Lyme disease, the elu- 
cidation of its causative agent, and the development 
of diagnostic tests and of treatments for the disease. 
| The disease was first recognized in North America 
when, around 1975, doctors at the Yale University 
School of Medicine noticed a new array of symptoms 
in patients from the area of Lyme, Connecticut. These 
symptoms consisted of a growing, ring-shaped rash 
(the bull’s-eye) at the site of a tick bite, and swollen 
joints. Sometimes the rash and the arthritis had ap- 
peared without the patient’s being aware of a tick bite. 
The author, Jonathan Edlow, M.D., is Vice-Chairman 
of the Department of Emergency Medicine, Beth Israel 
Deaconess Medical Center, and Assistant Professor 
of Medicine, Harvard Medical School. He himself 
describes the book as “a medical detective story” that 


_| flowed from a combination of his interest in infectious 


diseases, and the fact that relatives of his who had 


| moved to the Lyme area in the 1980s were stricken with 


symptoms of a bizarre arthritis, and/or rashes following 


_| tick bites. 


The author starts with a detailed and interesting 


j; account of how, in the 1970s, residents of the Lyme area 


were experiencing a baffling array of physical symp- 
toms, including rashes, unexplained neurological symp- 
toms and swollen joints, that eluded diagnosis by their 


"| family doctors. Then two women, who together with 
; their families and neighbours, had been ill for several 


years, were separately urged by their doctors to visit 
the Rheumatology Clinic at the Yale School of Medi- 
cine in New Haven. After hearing their stories, the doc- 
tors at the clinic realized that about thirty-five cases 


) : : ae 
clustered in the Lyme area were presenting similar 


! 


| Edited by Peter Kareiva and Simon A. Levin. Princeton 
University Press, 41 William Street, Princeton, New Jer- 
sey 08540. 427 pages. U.S.$53.00 Paper. 


The ecological impact of the economic system we 
subscribe to could be viewed as the largest of all pos- 


BOOK REVIEWS 


64] 


Bull’s Eye: Unraveling the Medical Mystery of Lyme Disease 


symptoms, and that an investigation into “what was 
happening at Lyme” was required. At the same time, 
U.S. Navy doctors at the nearby base in Groton, Con- 
necticut, observed several patients with ring-shaped 
rashes which spread over a large area. Their case report 
in The Journal of the American Medical Association 
(JAMA), was read by a family doctor at the eastern 
end of Long Island who had patients with tick bites 
surrounded by a rash, who thought that his patients 
might have the same disease. He realized it was similar 
to a disease that had been recognized in Europe since 
1910 as erythema migrans (EM). In the European 
reports, the disease was associated with tick bites and 
thought to have a spirochaete as the causative agent. 

Having set the scene, the author describes the iso- 
lation of this causative agent by Willy Burgdorfer at 
the Rocky Mountain Laboratories, U.S. Public Health 
Service, who demonstrated that a spirochaete found in 
the deer tick, Ixodes scapularis, caused Lyme disease. 
The organism was later named Borrelia burgdorferi 
in his honour. But this tick can carry other pathogens 
(e.g., Babesia spp.) causing a co-infection in some 
patients, complicating both diagnosis and treatment. 
He further describes how, over time, a definite schism 
arose between the two groups of doctors maintaining 
the differing viewpoints mentioned. Patient advocacy 
groups arose who mainly supported the “alternative” 
position and, in 1993, a U.S. Senate Committee held 
hearings on the subject. The author sums up the cur- 
rent position by saying that debate remains about the 
best way to diagnose the disease, the utility of long- 
term antibiotic treatment, and the safety of a vaccine. 

A drawback in the author’s style arises when he uses 
analogies to try to explain scientific concepts. One 
example is that of a five-page description (complete 
with 5 Tables!) of the separation of peaches, apples, 
oranges and nectarines by machine, in an attempt to 
explain the meaning of the terms sensitivity and speci- 
ficity in assessing the accuracy of laboratory tests. The 
result is confusion. 

Appendix A lists the symptoms of Lyme disease, 
Appendix B lists tick-borne diseases in humans and 
animals. The book does not have a bibliography but. 
following the appendices, gives a list of sources for 
each chapter. 

PEARL PETERKIN 


#801 — 240 Brittany Drive, Ottawa, Ontario K1K OR7 Canada 


The Importance of Species: Perspective on Expendability and Triage 


sible species removal experiments. Unfortunately for 
us, this experiment is unreplicatable—we only get one 
shot at it. From the mid-point things don’t look good. 
When the consequences of our personal actions and 
public policies are compared to the cataclysmic impact 


of an asteroid (we are in the midst of the sixth and 
greatest wave of extinctions on the planet after all) you 
know we're in trouble. Still, the overwhelming number 
of species facing imminent elimination has lead some 
to ask if we need to protect every one? Surely of the 
millions of species we “share” Earth with there must 
be a few that we could afford to lose? Especially con- 
sidering the expense and inconvenience of conserva- 
tion programs. 

“The importance of species” sets out to answer 
that question. Or at least to summarize the collective 
experiences of the ecological community in examin- 
ing the idea of species expendability. The initial impe- 
tus for this volume came from a symposium to hon- 
our the accomplishments of Robert Paine. Dr. Paine’s 
pioneering work on the importance of species began 
nearly 40 years ago. His investigations were based on 
a simple idea: if you want to understand the role of a 
particular species in a community, remove it and see 
what happens. He works primarily in rocky inter-tidal 
zones. It was there he discovered that some species 
are absolutely critical to that ecosystem: remove one 
of these “keystones” and the community collapses. 
This conclusion has since been reached by many other 
ecologists working in different systems, and the book 
provides a nice introduction to their work. However, 
while keystone species have been identified in a variety 
of settings, their redundant counterparts have been 
harder to locate. 

The strength of this book lies in the diversity of 
voices that converge on a few common conclusions. 
The most critical outcome of a great deal of research 
is the recognition that identifying truly redundant spe- 
cies is an extremely difficult and perhaps impossible 
task. Each paper presents new obstacles. Louda and 
Rand present a particularly nasty problem in the first 
chapter. Native thistles of the Great Plains may ini- 
tially appear to be expendable as a direct result of their 
being essential! They argue that the native thistles serve 
to inoculate the community against the invasion of their 
exotic relatives. The presence of the native thistles sup- 
ports populations of herbivorous insects which readily 
switch to feeding on exotic thistle species that have 
been introduced to the region. In areas where this has 
been most successful the exotic thistles have been al- 
most completely excluded from the community. Under 
these conditions the exotic thistles do not appear to 
pose a serious threat to the community, so the value 
of the native thistles is not readily apparent. 

More generally, the environmental context may 
determine the relative importance of a species in a 
community. A keystone predator at one location may 
have negligible influence at another (Chapter 2). Sim- 
ilarly, the influence of a species may vary greatly over 
time (Chapter 3). Think of a herbivorous insect that 
spends nine years in small, widely scattered populations 
of minor importance, only to erupt in the tenth year 
to devastate thousands of hectares of forest (and feed 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


millions of birds...). At a broader temporal scale, 
Stephen Palumbi (Chapter 15) notes that important 
ecological associations can evolve rapidly. A species 
that is redundant now may become an important com- 
ponent of its ecosystem in a few decades. The blink 
of an eye in evolutionary terms, but an eon measured 
in funding cycles. Taking a different approach Daniel 
Simberloff (Chapter 11) examines the impacts of the 
extinctions or near-extinctions of species one could 
reasonably assume to be important community mem- 
bers (i.e., American Chestnut, American Elm, Bison 
etc.). He reveals two related challenges: we have very 
little data to objectively assess the impacts of these 
losses, and there is no agreement on what constitutes 
a significant “ecosystem consequence”. 

As you would expect from a book with 28 authors 
the treatment is a little uneven. Given the emphasis on 
detailed natural history knowledge that many authors 
present as critical to the question, I was not convinced 
of the value of theoretical models in predicting which 
species are expendable. Shahid Naeem’s (Chapter 6) 
discussion of ecosystem reliability models starts from 
the assumption that species are interchangeable, effec- 
tively sidestepping the entire issue. 

The most troubling essay is perhaps the most impor- 
tant one in evaluating the concept of expendability. 
Mary Ruckelshaus, Paul McElhany, and Michael Ford 
collaborate in reviewing their efforts to conserve Pacific 
salmon species as part of the American National Marine 
Fisheries Service. Given that five of six of these pre- 
sumably economically important species are threatened 
or endangered, you might expect they would approach 
their task with the objective of maximizing the viability 
of the various stocks. On the contrary, they outline the 
process they followed in determining how many more 
populations they can afford to lose. This is a chilling 
glimpse of the possible future of conservation biology. 
The emphasis is not on how best to protect and enhance 
our biological heritage. Rather, conservation goals are 
set in a manner that minimizes their impact on eco- 
nomic objectives. 

Egbert Leigh’s contribution, “Social conflict, bio- 
logical ignorance, and trying to agree which species 
are expendable” (chapter 12) provides an insightful 
review of the entire debate. He identifies a critical need 
for basic natural history research if we are to have any 
hope of understanding the relative importance of indi- 
vidual species. More importantly, he reminds us “that 
social conflict, whether latent or blatant, is the biggest 
single obstacle to sound conservation. Suppose for a 
moment that we were Laplacean demons, able to assess 
the ecological consequences of each and every extinc- 
tion. Would this circumstance really cause society to 
close ranks behind conservation? As the world seems 
organized to destroy any sense of community among 
human beings, a greater knowledge of biology is not 
likely to make it easier for society to agree on conser- 
vation policy.” Leigh reinforces the value of natural 


2004 


| history in dealing with biological issues, while prop- 
| erly identifying the question of species expendability 
as part of a larger social problem. 

This is an important book, and deserves the critical 
attention of conservation biologists. In an age when we 
_ are pressured to apply a business model to all aspects 
of our society, it is important to examine the conse- 
| quences in a rigorous way. The essays in this book 


NEW TITLES 
+Available * Assigned 


Zoology 


*Common Birds of Ontario. By D. Sept. 2004. Calypso 
| Publishing, Box 1141, Sechelt, British Columbia VON 3A0. 
| 93 pages, Can. $12.95. 


) +Birding in Venezuela (revised edition). By M. Goodwin. 
| 2004 Lynx Edicions Barcelona Spain. 332 pages, U.S.$25.60 
| Paper. 


| Birds of the Middle East (Reprint of 1996 edition). By R. 


| Porter, S. Christensen and P. Schiermacker-Hansen. 2004. 


Princeton University Press, 41 William Street, Princeton, 
| New Jersey, USA, 08540-5237. 480 pages, U.S.$35. Paper, 
| U.S.$65 Cloth. 


| +Bumblebee Economics (2™ Edition). By B. Heinreich. 
| 2004. Harvard University Press, 79 Garden Street, Cambridge, 
Massachusetts 02138 USA. 245 pages, U.S.$19.95 Paper. 


_ *Damselflies of Alberta — Flying Neon Toothpicks in the 
| Grass. By John Acorn. 2004. University of Alberta Press, 
! Ring House 2, Edmonton, Alberta T6G 2E1 Can. $29.95. 


_ Experimental Approaches to Conservation Biology. Edit- 
_ ed by M. S. Gordon and S. M. Bartol 2004. University of 
California Press, 2120 Berkeley Way, Berkeley, California 
94704-1012. 358 pages, U.S.$75. 


*Frogs of Australia. By J. Turner. 2004. Pensoft Publishers. 
Acad. G. Bonchevy Str., Bl. 6, 1113 Sofia, Bulgaria. 276 pages, 
Eur 34.50 Cloth. 


*How the Earthquake Bird got its name and Other Tales 
of an Unbalanced Nature. By H. Shugart. 2005. Yale Uni- 
versity Press, P.O. Box 209040, New Haven, Connecticut 
06520-9040. 227 pages, U.S. $30. Cloth. 


Hummingbirds. By L. Aziz. 2004. Firefly Books Ltd., 3680 
Victoria Park Avenue, Toronto. 64 pages, Can. $9.95. Paper. 


*Mammals of Australia. By J. Turner. 2004. Pensoft Pub- 
lishers. Acad. G. Bonchev Str., Bl. 6, 1113 Sofia, Bulgaria. 
216 pages, Eur 38.50 Cloth. 


*The Peregrine Falcon Survey in Canada. Edited by U 
Banasch and G. Holroyd. 2004. Canadian Wildlife Service 
4999 — 98 Ave., Edmonton, Alberta T6X 2X3 Canada. No 
price available. 


Seabirds: a natural history. By A. Gaston. 2004. Yale Uni- 
versity Press, Box 209040, New Haven, Connecticut 06520. 
224 pages, U.S.$45.00. 


BOOK REVIEWS 


643 


allow us to do that. The motivation behind identifying 
redundant species is that it would allow us to maximize 
the bang per conservation buck. However, cost-benefit 
analysis is of very limited value when you cannot quan- 
tify either costs or benefits. 


TYLER SMITH 
155 Avenue Vanguard, Pointe Claire, Quebec H9R 3T4 Canada 


*Spiders of Australia. By T. Hawkeswood. 2004. Pensoft 
Publishers. Acad. G. Bonchev Str., Bl. 6, 1113 Sofia, Bulgaria. 
264 pages, Eur 34.95 Cloth. 


*Venomous Reptiles of the Western Hemisphere. By J. 
Campbell and W. Lamar. 2004. Cornell University Press, 
512 East State Street, Ithaca, New York 14850. 1032 pages 
(2 volumes), U.S.$149.95. 


*Whales and Dolphins of the World. By M. Simmonds. 
2004. The MIT Press, Five Cambridge Centre, Cambridge, 
Massachusetts 02142. 160 pages, U.S.$29.95 Cloth. 


Botany 


Atlas of Plants of Nunavik Villages. By M. Blondeau and 
C. Roy. 2004. Editions Multimondes, 930 rue Pouliot, Sainte- 
Foy, Quebec G1V 3N9. Can. $34.95. 


*Boreal Forest of Canada and Russia. By W. Pruitt and L. 
Baskin. 2004. Russian Academy of Science, Leniasky pr., 
33, Moscow V-71, Russia. 163 pages, Price not known. 


+Flower Chronicles (reprinted). By E. B. Hollingsworth. 
2004 (original 1958). University of Chicago Press, 1427 East 
60" Street, Chicago, Illinois 60637. 300 pages, U.S.$16. 


*Gathering Moss — A Natural and Cultural History of 
Mossess. By R. Kimmerer. 2004. Oregon State University 
Press, 102 Adams Hall, Corvallis, Oregan 97331. 176 pages, 
U.S.$17.95 Paper. 


Environment 


*Experimental Approches to Conservation Biology. Edit- 
ed by M. Gordon and S. Bariol 2004 University of Califor- 
nia Press. 343 pages, not illus. U.S.$ ? 


+Rendevous with the Wild — The Boreal Forest. Edited by 
J. Raffan (ed). 2004. BostonMills Press — Firefly Books Ltd.. 
3680 Victoria Park Avenue, Toronto, Ontario M2H 3K1 
188 pages, Can. $39.95 


Miscellaneous 


The New Amateur Naturalist. By Nick Baker .2004. Harper 
Collins HarperCollins Publishers Ltd., Bloor Street East, 
20" Floor, Toronto, Ontario M4W 1A8. 288 pages, £14.99 — 
approx. U.S.$38 Cloth. 


Wildlife Spectacles. By Russell A. Mittermeier et al. 2004. 
Conservation International 1919 M Street, NW Suite 600, 
Washington, DC 20036. 324 pages, £39.50 — approx. U.S.$74. 


News and Comment 


RENEW. 2004. Recovery of Nationally Endangered Wildlife in Canada. Annual Report (14) 


Contents (36 pages): Highlights of 2003-2004 [Of 160 
Endangered, 108 Threatened, and 21 Extirpated species on 
the November 2003 COSEWIC list: 206 species are wholly or 
partially covered by recovery plans (completed or in develop- 
ment); 117 species are included in ecosystem or multispecies 
recovery, to the extent that their range overlaps with the geo- 
graphic area; 53 show stable or increasing population trend. 
$30.7 million was expended on recovery (salaries + expenses): 
Employment was equivalent to about 142 salaried and 23 vol- 
unteer people working full-time; 258 organizations made finan- 
cial contributions; 348 organizations participated on recovery 
teams] — Report from the Co-Chairs — Figures [4] — Species 


Recovery — Responsible Jurisdictions — Financial Contributors 
— Funding per Target — Canadian Wildlife Directors Commit- 
tee [contacts for species at risk]. The report was produced 
by the Canadian Wildlife Service of Environment Canada in 
cooperation with the provinces, territories, Fisheries and 
Oceans Canada, Parks Canada Agency, Wildlife Management 
Boards and many individuals and non-government groups. 
Readers are cautioned (inside front cover): “This informa- 
tion is carefully reviewed but the Canadian Wildlife Service 
disclaims responsibility for the accuracy of the information 
contributed”. 


Calendar: Endangered Reptiles of Canada 2004 and 2005 


For 2004, 12 (one for each month) striking black-and- 
white drawings by various artists portray five turtles, 1 lizard 
and seven snakes whose surviving Canadian populations are 
designated as Endangered (Blue Racer), Threatened (Black 
Ratsnake, Eastern Foxsnake, Stinkpot, Eastern Ribbonsnake 
[Nova Scotia population], Blanding’s Turtle [Nova Scotia 
population], Eastern Massasauga), Special Concern (Wood 
Turtle, Five-lined Skink, Eastern Ribbonsnake [Ontario pop- 
ulations], Eastern Milksnake, Spotted Turtle), or Under Review 
(Prairie Rattlesnake, Blanding’s Turtle [Quebec, Ontario] 
populations). For each species there are brief remarks on ap- 
pearance, occurrence, habitat and risks to its survival. A 13% 
species, the Pacific Pond Turtle is featured on the cover as 
designated “extirpated” despite the lack of credible records 
that native populations ever existed in Canada with the spec- 
ulative remark “There is no biogeographic basis for the Pacific 
pond turtle’s range not to have extended in Canada, so cur- 
rent efforts to rebuild populations in Washington State may 
result in the return of the Pacific pond turtle to southwestern 
British Columbia”. The fine artwork was contributed by Joe 
Crowly (3), Kathryn Peiman (3), Mandi Eldridge (2), Kevin 
Kerr (2), Sonya Amin, Marisa Bonofiglio, and Sarah Ing- 
werson. 

In the 2005 version the black-and-white art is replaced by 
colour photography. The cover head of Western Rattlesnake 
(Crotalus oreganus) from British Colombia [sic] is followed 
by 7 snakes, 6 freshwater turtles, and one lizard each opposite 
a monthly calendar page covering December 2004 to Decem- 
ber 2005. Each has additional cameo shots of the same species 
and a paragraph that highlights its status and characteristics. 


Canadian Species at Risk November 2004 


The latest version of the booklet Canadian Species at Risk 
prepared after the November 2004 COSEWIC assessment 
meeting is available on-line in both PDF and HTML formats 


Species represented are Eastern Ratsnake (Elaphe obsoleta) 
[= Western Ratsnake Elaphe spiloides of one recent study], 
Eastern Foxsnake (Elaphe gloydi) [= Elaphe vulpina gloydi\, 
Wood Turtle (Glyptemys insculpta), Spiny Softshell Apalone 
spinifera), Stinkpot (Sternotherous odoratus), Massasauga 
(Sisturus catenatus), Eastern Hog-nosed Snake (Heterodon 
platirhinos), Five-lined Skink (Eumeces fasciatus), Blanding’s 
Turtle (Emydoidea blandingi), Spotted Turtle (Clemmys gut- 
tata), Racers (Coluber constrictor: 3 subspecies), Snapping 
Turtle (Chelydra serpentina), Great Basin Gopher Snake 
(Pituophis catenifer deserticola). The back of the calender 
features 3 species supposedly extirpated from Canada: Timber 
Rattlesnake (Crotalus horridus), Eastern Box Turtle (Terra- 
pene carpolina carolina) and Pacific Pond Turtle (Actinemys 
marmorata). Not mentioned 1s that the latter two were likely 
introductions to Canada. Two other forms are not illustrated 
but listed: Pacific Gophersnake (Pituophis catenifer catenifer) 
and Pigmy Short-horned Lizard (Phrynosoma douglasii). 

These calendars have been produced by “the Brooks lab” 
at the Department of Zoology, University of Guelph, Guelph, 
Ontario NIG 2W1 Canada; (519) 824-4120 extension 58360; 
e-mail brookslab@hotmail.com. The selections for both cal- 
endars are based on the latest status designations by the reptile 
subcommittee (for which Brooks in chair) of the Committee 
on the Status of Endangered Wildlife (COSEWIC). See website: 
www.cosewic.ga.ca. All proceeds from sale of calendars goes 
to support the Kawartha Turtle Trama Centre in Peterborough, 
Ontario, established in 2002 to treat injured native turtles and 
release them back in the wild (website: www.kawarthaturtle. 
org) and reptile conservation research. 


on the web site http://www.cosewic.qc.ca/eng/sct5/indexe.cfm 
Paper format of this publication will now be available only 
once a year, following the spring assessment meeting. 


644 


2004 


Number 65 (Focus on Australia), October 2004. Contents: 
| DAPTFE Seed Grants 2005 — The Impact of Habitat Modifica- 
| tion on the Striped Marsh Frog Limnodynastes peronii (Amy 
Gye) — Conservation Status of Australian Frogs (Jean-Marc 
Hero, Care Morrison, Graeme Gilespie, Dale Roberts. Paul 
Horner, David Newell, Ed Meyer, Keith McDonald, Frank 
Lemckert, Michhhhael Mahony, Michaelll Tyler, Will Osborne, 
Harry Hines, Steve Richards, Conrad Hoskin, Naomi Doak and 
Luke Shooo) — New Funds available from the DAPTE (Tim 
Halliday) — Alaska Conference Summary (Deborah Rudis — 
Froglog Shorts. 

| Number 66, December 2004. Contents: New Funds avail- 
_ able from the DAPTF (Tim Halliday & Don Church) — U.S. 
| Cricket Frog Symposium (Michael Lannoo and Ralph Grundel) 
_ — Press Release: Sunburt Frogs a Myth: Pond Scum offers 
| Natural Sunscreen (Larry Licht) — Global Amphibian Assess- 
/ ment (Simon Stuart, Janice Chanson, Neil Cox and Bruce 
| Young) — Atelopus mucubajiensis still survives in Andes of 
_ Venezuela. Prelimnary Report (Cesar L. Barrio-Ammmmoros, 


| December 2004 


Editor’s Note — Thank You from the Meeting Organizer — 
_ Overview of the Annual Meeting [Edmonton, Alberta, 24-27 
_ September 2004] — 2004 Silver Salamander Award Winners 
| [Laura Frits, Steve Brechtel] — 2004 Best Student Presenta- 
_ tion [Connie Browne] — Meeting Field Trip: Tyrrell Museum 

and Reptile World — Meeting Field Trip: Garter Snake Dens 


| Marine Turtle Newsletter (106) 


October 2004. 28 pages: EDITORIAL Conceptual Problems 
_ with the ICUN Red Listing Assessment for the Green Turtle: 
_ Move Over Raine Island — GUEST EDITORIAL: Sea Turtles, 
| Red Listing, and the Need for Regional Assessments — 
_ ARTICLES: The Trade of Marine Turtles in the Toliara region, 
_ Souty West Madagasgar — NOTES: Rat Eradication as Part of 
a Green Turtle (Chelonia mydas) Conservation Prograamme 
in Indonesia — Fibropapillomatosis and Multiple Fibromas 
in a Green Turtle from the South Coast of Pernambucoo State, 
Brazil — First Record of a Leatherback Turtle in Syria — MEETING 
REPORTS — IUCNMTSG UPDATE — ANNOUNCEMENTS — NEWS & 


NEWS AND COMMENT 


645 


| Froglog: Newsletter of the Declining Amphibian Populations Task Force (65, 66) 


Fundacion AndigennA — The Afelopus Initiative: Conserving 
Endangered Tropical Andeannn Amphibians (Ariadne Angulo, 
Paul Salaman and Jose Vicente Rodriguez — Book Reviews 
— Froglog Shorts. 

Froglog is the bi-monthly newsletter of the Declining 
Amphibian Populations Task Force of The World Conserva- 
tion Union (IUCN)/Species Survival Commission (SSC) and 
is supported by The Open University, The World Congress 
of Herpetology, and Arizona State University. These issues 
were edited by John WIlkinson and Jeanne McKay (65) and 
Jeanne McKay (66), Department of Biological Sciences, The 
Open University, Walton Hall, Milton Keynes, MK7 6AA, 
United Kingdom; e-mail: daptf@open.ac.uk. Funding for 
Froglog is underwritten by the Detroit Zoological Institute, 
P. O. Box 39, Royal Oak, Michigan 48068-0039, USA. Pub- 
lication is also supported by Peace Frogs www.peacefrogs.com 
and by RANA and the US National Science Foundation grants 
DEB-0130273. 


_ The Boreal Dip Net/L’Epuisette Boreale: Newsletter of the Canadian Amphibian and Reptile 
Conservation Network/Reseau Canadien de Conservation des Amphibiens et des Reptiles 9(1) 


— List of Presentations titles at 2004 Annual Meeting — 
Attendees of the 2004 Annual Meeting [55]. - Membership 
in CARCNET/RECCAR (contact Bruce Pauli, Canadian 
Wildlife Service, National Wildlife Research Centre, Carleton 
University, Raven Road, Ottawa, Ontario K1A 0H3. Web site: 
http://www.carcnet.ca/). 


LEGAL BRIEFS — RECENT PUBLICATIONS. 

The Marine Turtle Newsletter is edited by Brendan J. Godley 
and Annette C. Broderick, Marine Turtle Research Group, 
Centre for Ecology and Conservation, University of Exeter 
in Cornwall, Tremough Campus, Penryn TR10 9EZ United 
Kingdom; e-mail MTN @seaturtle.org; Fax +44 1392 263700. 
Subscriptions and donations towards the production of the 
MTN can be made online at <http://www.seaturtle.org/mtn/> 
or postal mail to Michael Coyne (online Editor) Marine Turtle 
Newsletter, 1 Southampton Place, Durham, North Carolina 
27705 USA (e-mail: mcoyne @seaturtle.org). 


WARS 1983-2002 Special Annual Report Wildlife Accident Reporting and Mitigation 


The British Columbia Ministry of Transportation has 
published as a CD this report on the Wildlife Accident 
Reporting System (WARS) which describes its work and 
achievements of the last 20 years as one of the pioneers and 
leaders in wildlife accident reporting and mitigation. 

The WARS system in used in British Columbia to: 

1) quantify the magnitude of wildlife-related vehicle colli- 

sions by species 

2) identify accident-prone locations and accident trends 

3) direct accident mitigation efforts (such as wildlife exclu- 

sion fencing and wildlife crossing structures) to problem 
locations 

4) evaluate the effectiveness of accident risk profiles for 

highway corridors 


5) develop species-specific accident risk profiles for high- 
way corridors 
6) advance mitigation efforts to protect rare and endangered 
species 
7) establish policies and strategies for accident issues and 
mitigation iniatives 
Copies of the CD can be obtained from Leonard Sielecki, 
R. P. Bio, MCIP, Engineering Branch, Environmental Manage- 
ment, British Columbia Ministry of Transportation, P.O. 
Box 9850, Station Provincial Government, Victoria British 
Columbia V8W 9T5 [location address: 4B — 940 Blanshard 
Street, Victoria] e-mail: leonard.sielecki@ gems9.gov.be.ca: 
telephone (250) 356-2255. 


The Ottawa Field-Naturalists’ Club Awards for 2003 


IRWIN M. BRopo, CHRISTINE HANRAHAN, BEVERLY MCBRIDE, and SHEILA THOMSON 


At the Club’s Annual Soirée, held on 24 April 2004, at St. 
Basil’s Church in Ottawa, awards were once again given to 
members, and one non-member, who distinguished them- 
selves by accomplishments in the field of natural history and 
conservation, or by extraordinary activity within the Club. 


Daniel Strickland — Honorary Member 

The Ottawa Field-Naturalists’ Club takes great pleasure in 
honouring Daniel Strickland, not only for his renowned stud- 
ies on Gray Jays, but also for the major role that he has played 
in fostering an appreciation and enjoyment of natural history 
by the general public. 

Dan earned his Bachelor of Science degree from the Uni- 
versity of Toronto, followed by his Master of Science from the 
University of Montreal, choosing as his thesis the life history 
of the Gray Jay, which he studied in La Verendrye Park in 
Quebec. 

For more than thirty years, Dan has been studying and docu- 
menting Gray Jay behaviour in Algonquin Park, including the 
dynamics of the Gray Jay family unit, its nesting behaviour, 
winter survival and food gathering strategies. He is recognized 
world-wide as an authority on Gray Jays, and has co-authored 
the account on Gray Jays in the prestigious series, The Birds 
of North America. 

In 1970, Dan became Chief Naturalist for Algonquin Park, 
a position he held until his retirement in 2001. Under his 
leadership, Algonquin Park’s nature interpretation program 
has become renowned continent-wide. Tens of thousands of 
park visitors each year enjoy the Park’s nature walks, public 


Colin Bowen — George McGee Service Award 2003 


When Colin Bowen first joined the Birds Committee 13 
years ago, committee members recognized a man who would 
do an excellent job at whatever he took on. Since then, he has 
put in countless hours of solid, thorough work to keep the 
OFNC’s many bird-related materials coherent and accessible. 

Colin is extremely well-organized. With his involvement, 
information management projects get done effectively. In the 
mid-90s the Birds Committee urgently needed to consolidate 
and properly archive over thirty years’ worth of bird records 
and publications that were scattered in homes around the region. 
Colin formed the backbone of a team that worked for almost 
three years to bring it all together. He continues to maintain the 
archives so that, now, anyone can easily find what they need. 

As secretary of the Bird Records Sub-committee, Colin has 
a major task on his hands. Sub-committee membership has 
changed many times since it first reconstituted itself in the 
mid-90s, but Colin has stayed. He receives all rare bird reports 
and prepares and redistributes them to committee members 
for review. He tracks all debates and decisions at meetings and 
prepares the sub-committee’s minutes and reports. On top of 
all that, he set up and maintains the extensive cross-referenced 
bird records database and photo collection. 

A visit to the OFNC’s website will reveal fine examples 
of Colin’s efforts to bring Ottawa bird resources to the Internet 


As is frequently the case, there isn’t a winner for every poten- 
tial award and, this year, the Anne Hanes Natural History 
Award was not given. The following citations for those who 
did receive an award, however, were read to the members 
and guests assembled for the event. 


wolf howls, and natural history slide presentations. In addition, | 


seventeen self-guided nature trails, dozens of natural history 
publications on subjects as diverse as butterflies, mushrooms, 
mammals and lichens, as well as a world-class interpretive 
centre, all had their genesis under Dan Strickland’s tenure as 
Chief Naturalist. In Algonquin Park’s lively and informative 
publication, The Raven, Dan shared his broad knowledge and 
his fascination with nature in more than 300 articles. The inter- 
pretive programs he developed have contributed immeasurably 
to the goal of fostering public understanding and appreciation 
of nature. 

Not the least of Dan Strickland’s legacies is his impact on 
a whole generation of young park naturalists, trained under 
his guidance, who have moved on to new parks or to new 
fields of endeavour, profoundly influenced by Dan’s insight 
and his enthusiasm for natural history. In recognition of his 
contributions to our knowledge of the natural history of Gray 
Jay, as well as his outstanding career in the field of nature 
interpretation and public education, it is a pleasure to confer 
Honorary Membership in the Ottawa Field-Naturalists’ Club 
on Dan Strickland. 


and therefore to a broader community. Visitors can see the on- 
line. tabular seasonal Ottawa bird checklist, the beautifully 
tabulated Christmas Bird Count results or the often-consulted 
Ottawa birding site guide, complete with maps. 

Colin has volunteered for the Peregrine Falcon Watch and 
the Ontario Breeding Bird Atlas. He has helped coordinate 
many Christmas and Fall Bird Counts. When out of town bird- 
ers contact the club looking for birding information Colin is 
one of the generous volunteers who offer hours of their time 
to take them to our fine birding spots. 

Colin does so much work for the Birds and Bird Records 
committees that we wonder how he finds time to do anything 
else... but he does. He is a devoted and highly-valued volun- 
teer at the Canadian Museum of Nature and, together with his 
wife, Pat, were the featured volunteers in the museum’s recent 
newsletter. He is a grandfather, an amateur genealogist, a trav- 
eller, and is taking up woodworking. He can also fix or build 
just about anything. 

The OFNC thanks Colin for his tremendous contribution, 
and for retiring from his job with Bell Canada so he would 
have more time to come and work with us. He has truly never 
stopped, and we are proud to present him with this year’s 
George McGee Service Award. 


646 


2004 


CLUB AWARDS 


647 


Daniel Brunton — Conservation Award for Members, 2003 


The recipient of the 2003 Conservation Award for Mem- 
bers, Daniel Brunton, is an individual well known to mem- 
bers of the Ottawa Field-Naturalists’ Club through his active 
role in Club affairs for over 20 years, including a term as 
President. Indeed, his name is familiar to many residents of 
the Ottawa region because he is frequently consulted and 
quoted as an expert naturalist and conservationist by the 
Ottawa media. Dan has been an ecological consultant since 
1979. The award he is receiving today, however, has to do 
with only one of his many activities on behalf of wildlife and 
wild spaces in the region, namely, his role in the establish- 
ment of the RiverKEEPER project for the Ottawa River. Dan 
has done almost 200 studies along the river, both as a profes- 
sional and amateur naturalist, making him eminently qualified 
to take this initiative. 

The problems associated with keeping track of the posi- 
tive and negative activities that impact the well-being of the 
Ottawa River demanded a new approach at coordination. 
With federal, two provincial, and dozens of municipal juris- 
dictions coming into play, and the high level of pollution af- 
fecting this crucial waterway, an initiative with vision was 
needed. Dan Brunton, together with some like-minded con- 
servationists, particularly George Brown, a Law professor at 


the University of Ottawa, and John Almstedt, got together to 
tackle the problem and recommended that the United States- 
based RiverKEEPER Alliance would provide the best frame- 
work within which to proceed. In the spring of 2001, a new 
Canadian chapter of RiverKEEPER was established with Dan 
as President. Its mission was to oversee the activities and 
legislation affecting the health of the river, and to report to the 
public. This would be done by hiring a full time KEEPER. 
Dan worked tirelessly to secure funding for the project and, 
after much work, succeeded in getting a three-year Trillium 
grant. In 2002, Laura Van Loon was appointed River KEEPER, 
launching the active phase of the project, and, in 2003, Dan 
was able to step down as President and allow others to carry 
on. 

As aresult of Dan Brunton’s efforts and vision, the public 
now has a full-time ombudsman to look after the river, advo- 
cating compliance with environmental laws, responding to 
citizen complaints, identifying problems that affect the river, 
and suggesting remedies. We are all beneficiaries of these 
efforts and have Dan Brunton to thank for helping to get the 
project started. The OFNC is therefore very pleased to pres- 
ent him with this year’s Conservation Award for Members. 


Friends of the Jock River — Conservation Award for Non-Member, 2003 


The OFNC Conservation Award, non-member, is given to 
a group or individual who has made an outstanding contribu- 
tion in the cause of natural history conservation in the Ottawa 
Valley, with particular emphasis on activities within the 
Ottawa District. 

The Jock River originates in the Goodwood Marsh approx- 
imately 50 kilometers west of Ottawa and empties into the 
Rideau River after winding through fields, villages, and the 
provincially significant Richmond Fen. Many species of fish, 
birds, mammals and other wildlife depend on this waterway 
for food, breeding sites and shelter. Over the years large areas 
of riparian vegetation have been destroyed causing severe 
silting of the river, which in turn has had a serious impact on 
flora and fauna of the river and its shoreline. The Friends of 
the Jock River, a non-profit, volunteer environmental organ- 
ization, was formed to look at ways of protecting, restoring, 
and enhancing the Jock River and its watershed. 

Working with local landowners, government agencies, 
recreational users and businesses, the Friends have engaged 
in numerous projects aimed at restoring the health of the river’s 
ecosystem. The number and types of work they have under- 
taken is truly impressive. 

The Friends have been planting native trees and shrubs at 
various sites along the river and this is helping restore sections 
of the river’s shoreline to a more natural forested state. One 
example of their work is the Twin Elm restoration project. 
This project combines restoration planting with fencing of 


Pearl Peterkin -OFNC Member of the Year, 2003 


This award is given to a member judged to have made sig- 
nificant contributions to the Club during the previous year. 
Looking back over the previous recipients of this award, it is 
illuminating to see the many and varied services performed 
for the Club. From rejuvenating committees, leading an ex- 
ceptional number of walks, editing Trail & Landscape, web 
site design and maintenance, the list is endless. But one thing 
is clear: joining the ranks of these hard-working, dedicated 
souls is no small thing. 


livestock from the riverbank and installation of a nosepump 
to meet their needs. Another important project undertaken 
by the group is The Rideau Valley Conservation Authority 
(RVCA) Stream Corridor Riparian Vegetation Survey. With 
training provided by the Conservation Authority, they col- 
lected data according to OMNR stream survey protocols to 
determine the ecological health of the river and its shoreline 
habitat. These data will become part of the RVCA database, 
which, in turn, will be made available to the Friends on CD 
to help them plan further riparian restoration projects. 

The Friends of the Jock River are involved with issues 
affecting the entire Jock River Watershed, thus they have been 
monitoring the ecology of the Richmond Conservation Area 
and advocating environmentally sound sewage treatment in 
Munster. 

The group is working on detailed guidelines for all future 
projects, including erosion control, buffering run-off, and en- 
hancing biodiversity in the watershed. They produce a quar- 
terly newsletter, hold regular meetings to which the public 
is invited, and maintain an informative website. 

For their dedicated, inspired, hard-working and thought- 
ful approach to protecting the health of the Jock River and 
its watershed, the OFNC is very pleased to give the Friends 
of the Jock River this Conservation Award. 

[Brian Finch, the present President; Eric Snyder, the 
Secretary; and Neil Barrington, the Past President, accepted 
the award on behalf of the Friends of the Jock River.] 


About ten years ago, it was decided to comb through back 
issues of Trail & Landscape and select those issues contain- 
ing articles of significant interest and value (for example, 
special issues on Butterflies, Reptiles and Amphibians, etc.) 
and retain a stockpile of these issues to sell. Since that orig- 
inal exercise, however, many more articles of interest have 
been published and Pearl Peterkin decided it was time to go 
through another selection process. She brought her sugges- 
tion to the Education and Publicity Committee who heartily 


648 


endorsed it. A suggestion was made to bundle together all 
issues on a particular topic and sell them as sets. This meant 
dealing with the entire 35 year run of Trail & Landscape, find- 
ing all the relevant issues, combining them, creating multiple 
sets, pricing them, finding adequate storage space at the Fletch- 
er Wildlife Garden centre and, of course, transporting the fin- 
ished products to the monthly meetings. During the past year, 
Pearl was often found ensconced in the back room of the 
Fletcher Centre, surrounded by stacks of Trail & Landscape, 
creating order out of chaos. She has also spent a lot of time 
there organizing the Education and Publicity Committee’s 
items so that they are more easily and quickly found. 

But it wasn’t only during 2003 that Pearl was active in Club 
affairs. Over the 15 years that Pearl has been an OFNC mem- 
ber, she has been involved with several club committees. In 
the 1990s she volunteered for three years with the Fletcher 
Wildlife Garden, becoming one of the stalwarts of this ener- 
getic group. In 2000 she took over as Chair of the Education 
and Publicity Committee at a time when the committee was 
at a low ebb with no leader. During her three years as Chair 
she successfully rejuvenated the committee, rounding up new 
members, finding judges for the Science Fair (which members 
of this Committee have been involved with for some time), 
finding volunteers to staff displays at various events, and do- 
ing all the other myriad jobs that keep a committee running 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


smoothly. Once the committee was up and running smoothly, 
Pearl decided to step down from that role and concentrate on 
other aspects of Education and Publicity committee work. 

Many people probably recognize Pearl, even if they don’t 
know her name, because she is the friendly face behind the 
OFNC sales table at the monthly meetings. It requires a seri- 
ous commitment to attend every meeting, and, when people 
rely on you to be there, the responsibility is that much greater. 
Each month Pearl has to gather together the various sale items 
and, with the help of some other volunteers, transport them 
to and from the meeting. She also has to keep on top of new 
items (such as the new Bird Checklist), and make sure the 
stock of regular items is in good supply. This dedication has 
paid off, however, because during the last year the sales table 
has become a focus of much interest and activity. 

In her spare time (what spare time, you may ask), Pearl 
is an enthusiastic and long time member of the Rideau Trail 
Association. She also volunteers for Bird Studies Canada’s 
Loon Survey. 

The OFNC is fortunate to have such enthusiastic and com- 
mitted volunteers as Pearl Peterkin. For hard work on the Trail 
& Landscape project as well as all her involvement with the 
Education and Publicity Committee and her dedication to 
the club, we are pleased to give Pearl Peterkin the Member 
of the Year award for 2003. 


. Errata for The Canadian Field Naturalist 118(3) 


| Inside front cover bottom: and marine algae, and, still lower, the algae take over. This 
Cover. Intertidal zone on rugged rock headland of the Queen — caption and photo (the latter in colour) have previously 
Charlotte Islands, British Columbia. Each dominant species —_ appeared as Figure 64, page 77, in Lichens of North America 
creates a distinct zone. The white lichen is mainly Coccotrema _ by I. M. Brodo, S. D. Sharnoff, and S. Sharnoff. 2001. Yale 
maritimum and the black one is almost entirely Verrucaria University Press, New Haven Connecticut. See article by 
maura. Below the Verrucaria is a zone bare of both lichens — Brodo and Sloan on lichen zonation, pages 405-424. 


On page 408, Figure 2, the islands were omitted. 


132°W 131°W 


53°N 


53°N 


18 Limestone 
Island 


Moresby 
Island 


Ep 


Gwaii Haanas 
National Park Reserve 
and Haida Heritage Site 
and proposed 
National Marine 
Conservation Area 


52°N 


17 
Treat Bay 


52°N 


50 Kilometres 


132°W 131°W 


FiGuRE 2. Map of the Gwaii Haanas area showing Limestone Island (location 18) to the north and Treat Bay (location 17) 
to the south of the insets. 


649 


Index to Volume 118 
Compiled by Leslie Cody 


Abies balsamea, 46,95,124,251,268,274,361,388,401,510,530 
lasiocarpa, 191,355,584 
Abrostola urentis, 541 
Acanthopteroctetes bimaculata, 532 
Acanthopteroctetidae, 532 
Acasis viridata, 539 
Acer, 52 
negundo, 309 
rubrum, 41,96,136,136,251,277,361,508 
saccharinum, 41,124,251 
saccharum, 361,507 
spicatum, 362 
Achigan a grande bouche, 523 
a petite bouche, 523 
Achillea millefolium, 166 
Achnatherum richardsonii, 564 
Acipenser fulvescens, 523 
Acipenseridae, 523 
Acleris albicomana, 534 
britannia, 534 
celiana, 534 
emargana, 535 
forbesana, 534 
logiana, 534 
nigrolinea, 535 
obligatoria, 534 
schalleriana, 534 
variana, 534 
Acorn, J.H., Review by, 468 
Acossus centerensis, 534 
populi, 534 
Acrobasis betulella, 536 
Acrocercops sp., 532 
astericola, 532 
Acronicta fragilis, 541 
grisea, 541 
impleta, 541 
impressa, 541 
innotata, 541 
vulpina, 541 
Acronictinae, 541 
Actaea X ludovici, 612 
Actitis macularia, 444 
Adalia bipunctata, 312 
Adela purpurea, 532 
Adelidae, 532 
Adelolecia koalensis, 406 
Aechmophorus spp., 210 
Aepyceros melampus, 185 
Aesculus pavia, 612 
Aethalura intertexta, 537 
Aethes promptana, 535 
promptana, 547 
Aethusa cynapium, 613 
Agabus sp., 431 
arcticus, 431 


confines, 431 
thomsoni, 431 
Agelaius phoeniceus, 201 
Agelaius phoeniceus, Staging in Eastern South Dakota, Spring 
Dispersal Patterns of Red-winged Blackbirds, 201 
Aglais milberti, 537 
Agonopterix gelidella, 533 
Agriphila ruricolella, 536 
vulgivagella, 536 
Agropyron spp., 400,584 
cristatum, 300 
pectiniforme, 560 
repens, 300 
sibiricum, 560 
Agrostis sp., 169 
filiculmis, 558 
gigantea, 560 
Aira praecox, 177 
Aix sponsa, 131,230 
Alaska, Changes in Loon (Gavia spp.) and Red-necked Grebe 
(Podiceps grisegena) Populations in the Lower Mata- 
nuska-Susitna Valley, 210 
Alaska, Interactions of Brown Bears, Ursus arctos, and Gray 
Wolves, Canis lupus, at Katmai National Park and Pre- 
serve, 247 
Alaska, Seed Dispersal by Brown Bears, Ursus arctos, in 
Southeastern, 499 
Alaska, The Heather Vole, Genus Phenacomys, in, 438 
Alberta: Distribution and Status, The Barred Owl, Strix varia 
in, 215 
Alberta, First Record of Mountain Lions, Puma concolor, in — 
Elk Island National Park, 605 
Alberta, Including New Provincial Records, Moths and But- 
terflies (Lepidoptera) of the Boreal Mixedwood For- 
est near Lac La Biche, 530 
Alberta, Winter Habitat Use by Wolves, Canis lupus, in Rela- — | 
tion to Forest Harvesting in West-central, 368 | 
Alca torda, 258 
Alces alces, 40,57,61,96,134,185,248,368,.606 
Alder, 46,247,268,401 
Speckled, 253 
Alewife, 319 
Alfalfa, 361 
Algae, 320 
Allium schoenoprasum ssp. sibiricum, 558 
Alnus sp., 46,247,401 
crispa, 268 i 
incana, 253 
incana ssp. rugosa, 136 
incanatable, 511 
rugosa, 268 
Alopecurus carolinianus, 170 
Alopex lagopus, 602 
Alopex lagopus, den by a Polar Bear, Ursus maritimus, Exca- 
vation of an Arctic Fox, 602 
Alosa pseudoharengus, 319 


| 
| 
| 


650 | 


4 


| 


1 


— 2004 


sapidissima, 77 

_ Alucita lalannei, 536 

| Alucitidae, 536 

Alyssum americanum, 570 

| obovatum, 558 

| Amandinea coniops, 411 
Amaranth, Purplish, 613 
Amaranthus blitum, 613 
Ambloplites rupestris, 523 

_ Amblyptilia pica, 536 

| Ambrosia x helenae, 613 

Ameiurus nebulosus, 523 

Amelanchier sp., 196 

alnifolia, 558 


| Amirault, D.L., 448 
i Amirault, D.L., J. Kierstead, P. MacDonald and L. MacDon- 


nell. Sequential Polyandry in Piping Plover, Chara- 
drius melodus, Nesting in Eastern Canada, 444 
| Ammodytes sp., 262 


| Ammophila breviligulata, 444 


_ Amphipacifica: Journal of Aquatic Systematic Biology 3(4), 
294 

| Amphipod, 77 

_ Amphipoea americana, 541 

| Ampithoe vallida, 79 

| Anabeana sp., 320 

| Anacampsis conclusella, 534 

| Anaplectoides prasina, 543 

| pressus, 543 


| Anas crecca, 131 


| platyrhynchos, 130 

| rubripes, 129,268,604 

_ Anas rubripes, Broods, Extraordinary Size and Survival of 
| American Black Duck, 129 

_ Anathix puta, 542 

_ Anatis labiculata, 312 


_ Ancylis comptana, 535 
i diminutana, 535 
fuscociliana, 535 


laciniana, 535 

subaequana, 535 
Andres, B.A., 210 
Andromeda polifolia, 591 


_ Andropogon gerardii, 342 


Andropolia contacta, 542 

Anguilla rostrata, 523 

Anguille d’ Amérique, 523 

Anguillidae, 523 

Annotated List of the Arctic Marine Fishes of Canada, 156 
Anser caerulescens, 602 

Antelope, Roan, 185 


| Anthoxanthum odoratum, 177 


Anthriscus caucalis, 613 
Anthyllis vulneraria, 613 


| Anticlea multiferata, 539 


vasiliata, 539 
Apamea cogitata, 541 

commoda, 541 
Aplectoides condita, 543 
Aplodinotus grunniens, 523 
Apotomis capreana, 535 

deceptana, 535 

infida, 535 

removana, 535 
Apple, 299,361 


INDEX TO VOLUME 118 


65] 


Apple, Crab, 125 
Apps, C.D., 191 
Arabis caucasica, 558 
holboellii var. retrofracta, 558 
holboellii var. secunda, 558 
Aralia nudicaulis, 46,365 
Arbutus menziesii, 174 
Arbutus, 174 
Arceuthobium americanum, 595 
douglasti, 595 
laricis, 595 
Arceuthobium americanum, on Jack Pine, Pinus banksiana, 
Growth in Manitoba, Effects of Lodgepole Pine Dwarf 
Mistletoe, 595 
Archilochus colubris, 85 
Archilochus colubris, Entanglements in Burdock, Arctium 
spp., at Delta Marsh, Manitoba, Ruby-throated Hum- 
mingbird, 85 
Archips argyrospila, 535 
Arctiidae, 540 
Arctium spp., 85,253,611 
lappa, 85 
minus, 85 
tomentosum, 85 
Arctium spp., at Delta Marsh, Manitoba, Ruby-throated Hum- 
mingbird, Archilochus colubris, Entanglements in 
Burdock, 85 
Ardea herodias, 231 
Argyresthia sp., 532 
abies, 532 
conjugella, 532 
goedartella, 532 
oreasella, 532 
pygmaeella, 532 
Aristida basiramea, 276 
Aristida basiramea Engelm. ex Vasey: A New Addition to 
the Flora of Quebec, Forked Three-awned Grass, 276 
Armoracia rusticana, 558 
Arrow-grass, Marsh, 561 
Arrowhead, 318 
Artemisia tridentata, 300,584 
Arthonia phaeobaea, 410 
Arthurs, W.R., Review by, 475 
Asaphocrita sp., 533 
Ascaphus truei, 447 
Ascaphus truei, American Dipper, Cinclus mexicanus, Preys 
Upon Larval Tailed Frogs, 446 
Ash, 253 
Black, 310 
Green, 309 
Mountain, 362 
White, 361,508 
Aspen, Quaking, 211,584 
Trembling, 46,62,96,218,251,268,274,277,316,342,355 
361,370,388,5 15,530 
Asphodel, Northern False, 567 
Aspicilia caesiocinerea, 411 
supertegens, 424 
Aspidotis densa, 166 
Aster spp., 251 
alpinus ssp. vierhapperi, 558 
falcatus, 558 
Aster, 251 
Alpine, 577 
Gulf of Saint Lawrence, 105 


652 


Western Heath, 577 
Aster, Symphyotrichum laurentianum, (Fernald) Nesom, Esti- 
mation of Seed Bank and Seed Viability of the Gulf 
of Saint Lawrence, 105 
Atkinson, J., Review by, 283 
Atriplex alaskensis, 558 
patula, 560 
patula sensu, 568 
patula var. alaskensis, 568 
subspicata, 559 
Auklet, Cassin’s, 616 
Rhinoceros, 615 
Auklet, Cerorhinca monocerata, Fledglings Fly to the Sea 
from Their Natal Burrows?, Do Rhinoceros, 615 
Austrocedrus chilensis, 132 
Autographa ampla, 541 
bimaculata, 541 
mappa, 541 
rubida, 541 
Avena sativa, 361,560 
Avens, Yellow, 573 
Awlwort, 572 
Aythya collards, 268 
marila, 268 


Bacidia alaskensis, 412 
Bacidina inundata, 424 
Baeus sp., 122 
Baeus sp. and Gelis sp., in the Egg Sacs of the Wolf Spiders 
Pardosa moesta and Pardosa sternalis (Araneae, Lyco- 
sidae) in Southeastern Idaho, Occurrence of Parasi- 
toid Wasps, 122 
Baillie, S.M., C.D. Wilkerson and T.L. Newbury. “Ashkui” 
Vernal Ice-cover Phenomena and Their Ecological 
Role in Southern Labrador, 267 
Balanus, 406 
Ball, P.W., 266 
Baneberry, Hybrid, 612 
Barbotte brune, 523 
Barbue de riviére, 523 
Barker, W., Review by, 474 
Barley, 361 
Barnacle, 406 
Bartier, P.M., 77 
Basilarchia arthemis, 537 
Bass, Largemouth, 113 
Basswood, 310 
American, 253 
Batrachedra praeangusta, 533 
Batrachedridae, 533 
Bear, 134,443 
Barren-ground Grizzly, 239 
Black, 41,96,370 
Brown, 247,499 
Grizzly, 117,239,247,370,603 
Polar, 395,602 
Bear, Ursus maritimus, Excavation of an Arctic Fox, Alopex 
lagopus, den by a Polar, 602 
Bears, H. Parasite Prevalence in Dark-eyed Juncos, Junco 
hyemalis, Breeding at Different Elevations, 235 
Bears, Ursus arctos, and Gray Wolves, Canis lupus, at Katmai 
National Park and Preserve, Alaska, Interactions of 
Brown, 247 
Bears, Ursus arctos, in Southeastern Alaska, Seed Dispersal 
by Brown, 499 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Bears, Ursus arctos, in the Central Canadian Arctic, Move- 
ments of Subadult Male Grizzly, 239 
Bears, Ursus maritimus, in the Alaskan Beaufort, Chukchi, 
and Northern Bering Seas, Observations of Habitat 
Use by Polar, 395 
Beaver, 40,129,248,267,606 
Eurasian, 434 
Beaver, Castor fiber, Collapsing Burrow Causes Death of a 
Eurasian, 434 
Bedford, R.E., 626 
Beech, American, 41,361,507 
Beetle, 388 
Carabid Intertidal, 79 
Colorado Potato, 226 
Convergent Lady, 309 
Lady, 309 
Ladybird, 225 
Parenthesis Lady, 309 
Seven-spotted Lady, 309 
Thirteen-spotted Lady, 309 
Transverse, 309 
Water, 425 
Beetles (Coccinellidae) in Overwintering Sites in Manitoba*, 
Density and Survival of Lady, 309 
Beetles (Coleoptera: Adephaga: Dytiscidae, Gyrinidae) Col- 
lected Along the Horton and Thelon Rivers in the Arc- 
tic Central Barrens of Canada, Predaceous Water, 425 
Beetles (Coleoptera: Coccinellidae) in Prince Edward Island, 
Use of Eelgrass, Zostera marina, Wrack by Three 
Species of Ladybird, 225 
Bennett, B., 558 
Bentgrass, 169 
Berberis spp., 132 
Beringer, J., 185 
Berube, D.K.A., 386 
Besma quercivoraria, 538 
Betula, 52 
alleghaniensis, 41,361,508 
glandulosa, 591 
papyrifera, 41,46,95,136,211,251,268,274,355,388,510, 
530 
populifolia, 277 
Bibarrambla allenella, 533 
Bidens sp., 180 
Bigler, L.L., 65 
Birch, Dwarf, 591 
Gray, 277 
Paper, 211,268,355 
White, 41,46,95,251,274,388,515,530 
Yellow, 41,361,507 
Bird rape, 571 
Bison bison, 115,606 
Bison, 115,606 
Bison, Bison bison, Toward Elk, Cervus elaphus, and Wolves, 
Canis lupus, Unusual Behavior by, 115 
Bison bison, Toward Elk, Cervus elaphus, and Wolves, Canis 
lupus, Unusual Behavior by Bison, 115 
Biston betularia, 537 
Bistort, Alpine, 591 
Bittercress, Alpine, 571 
Narrow-leaved, 613 
Blackbird, Red-winged, 201 
Yellow-headed, 207 
Blackbirds, Agelaius phoeniceus, Staging in Eastern South 
Dakota, Spring Dispersal Patterns of Red-winged, 201 


i 


— 2004 


Bladderwort, Lesser, 576 


| Blarina brevicauda, 402 


Blastodacna curvilineella, 533 


_ Bleptina caradrinalis, 540 


Blueberry, 251 
Lowbush, 195 
Tall, 575 
Velvet Leaf, 195 


| Blueberry, Vaccinium angustifolium, and Velvet Leaf Blue- 


berry, Vaccinium myrtilloides, Production in Eastern 
Ontario, Effect of Fire Intensity and Depth of Burn 
on Lowbush, 195 

Blueberry, Vaccinium myrtilloides, Production in Eastern 
Ontario, Effect of Fire Intensity and Depth of Burn 
on Lowbush Blueberry, Vaccinium angustifolium, 
and Velvet Leaf, 195 


_ Bluegill, 318 
| Bluegrass, 300 


Kentucky, 176 
Sandberg, 564 
Bluestem, Big, 342 
Bobcat, 40 
Bog-rosemary, Dwarf, 591 


| Bombycilla cedrorum, 232 


Bonasa umbellus, 41,232 
_ Bondia crescentella, 536 
_ Boreal Dip Net/L’Epuisette Boreal: Newsletter of the Canadian 
| Amphibian and Reptile Conservation Network — Re- 
seau Canadien de Conservation des Amphbiens et des 
| Reptiles, The, 155,484,645 
- Botrychium sp., 365 


| Bousfield, E.L, 626 
| Bowen, K.D. Frequency of Tail Breakage of the Northern 


Watersnake, Nerodia sipedon sipedon, 435 


| Bowerbankia gracilis, 79 


Brachylomia algens, 542 
discinigra, 542 


, Brachythecium spp., 253 
| Bracken, 253 


Branta canadensis, 268,602 
Brassica rapa, 560 
Braya glabella, 550 
humilis, 550 
pilosa, 550 
purpurascens, 550 
thorild-wulffii, 550 
Braya, Pilose, 550 
Braya, Braya pilosa Hooker (Cruciferae; Brassicaceae), an 
Enigmatic Endemic of Arctic Canada, Pilose, 550 
Braya pilosa Hooker (Cruciferae; Brassicaceae), an Enigmat- 
ic Endemic of Arctic Canada, Pilose Braya, 550 
Brisson, J. Forked Three-awned Grass, Aristida basiramea 
Engelm. ex Vasey: A New Addition to the Flora of 
Quebec, 276 
Bristlegrass, Green, 564 
British Columbia, Bird Communities of the Garry Oak Habitat 
in Southwestern, 376 
British Columbia, Introduced Marine Species in the Haida 
Gwaii (Queen Charlotte Islands) Region, 77 
British Columbia, Lichen Zonation on Coastal Rocks in Gwati 
Haanas National Park Reserve, Haida Gwaii (Queen 
Charlotte Islands), 405 
British Columbia, 1972-2002, Sea Otter, Enhydra lutris, Sight- 
ings off Haida Gwaii/Queen Charlotte Islands, 270 


INDEX TO VOLUME 118 


653 


British Columbia, Observations of Above-Surface Littoral For- 
aging in Two Sea Ducks, Barrow’s Goldeneye, Buceph- 
ala islandica, and Surf Scoter, Melanitta perspicillata, 
in Coastal Southwestern, 264 
British Columbia waters: a reassessment, Status of marine 
turtles in, 72 
Brodo, I.M. and N.A. Sloan. Lichen Zonation on Coastal 
Rocks in Gwaii Haanas National Park Reserve, Haida 
Gwaii (Queen Charlotte Islands), British Columbia, 
405 
Brome, Barren, 176 
California, 174 
Downy, 300 
Rip-gut, 174 
Smooth, 300,342,562 
Soft, 174 

Bromus spp., 175 
carinatus, 174 
ciliatus, 559 
hordeaceus, 174 
inermis, 300,342,560 
rigidus, 174 
sterilis, 176 
tectorum, 174,300 

Brook, R.K., 602 

Broom, Scotch, 177 

Brunton, D.F. Origins and History of The Ottawa Field- 
Naturalists’ Club, 1 

Brush, Big Sage, 584 

Bryotropha sp., 534 

Bryozoan, 79 

Bubo virginianus, 222,273 

Bucculatricidae, 532 

Bucculatrix sp., 532 

canadensisella, 532 

Bucephala clangula, 267,604 

islandica, 264 

Bucephala islandica, and Surf Scoter, Melanitta perspicillata, 
in Coastal Southwestern British Columbia, Observa- 
tions of Above-Surface Littoral Foraging in Two Sea 
Ducks, Barrow’s Goldeneye, 264 

Buckeye, Red, 612 

Budworm, Spruce, 361 

Bulrush, 318 

Bunchberry, 46 

Bunting, Indigo, 232 

Bur-chervil, 613 

Burdock, 85,253,611 

Common, 85 

Burdock, Arctium spp., at Delta Marsh, Manitoba, Ruby- 
throated Hummingbird, Archilochus colubris, Entan- 
glements in, 85 

Burnett, D., Review by, 151 

Burrows, F.G.M., 95 

Buteo lagopus, 268 

Buttercup, Western, 570 

Butterflies (Lepidoptera) of the Boreal Mixedwood Forest 
near Lac La Biche, Alberta, Including New Provin- 
cial Records, Moths and, 530 

Butterfly, 388 

Buttons, Brass, 169 


Cabera erythemaria, 538 
variolaria, 538 
Caenurgina crassiuscula, 541 


654 


Calafate, 132 
Calamagrostis canadensis, 591 
Calendar: Endangered Reptiles of Canada 2004 and 2005, 
644 
Callitriche anceps, 559 
Callizzia amorata, 539 
Caloplaca, 420 
chlorina, 424 
citrina, 411 
exsecuta, 420 
flavogranulosa, 411 
inconspecta, 406 
litoricola, 406 
marina, 420 
rosei, 411 
verruculifera, 411 
Caloptilia alnivorella, 532 
anthobaphes, 532 
betulivora, 532 
canadensisella, 532 
coroniella, 532 
stigmatella, 532 
Caloreas occidentella, 536 
Camas, Common, 175 
Great, 175 
Camassia leichtlinii, 175 
quamash, 175 
Camelina sativa, 558 
Campaea perlata, 538 
Campbell, B., 521 
Campion, Bladder, 569 
Canachites canadensis, 41 
Canada, Conservation Evaluation of Howell’s Triteleia, Tri- 
teleia howellii, an Endangered Lily in, 174 
Canada, Conservation Evaluation of Lemmon’s Holly Fern, 
Polystichum lemmonii, a Threatened Fern in, 164 
Canada, Conservation Evaluation of Small-flowered Lipo- 
carpha, Lipocarpha micrantha (Cyperaceae), in, 179 
Canada, Conservation Evaluation of the Pacific Population 
of Tall Woolly-heads, Psilocarphus elatior, an Endan- 
gered Herb in, 169 
Canada, Hybridization between a Green Turtle, Chelonia 
mydas, and Loggerhead Turtle, Caretta caretta, and 
the First Record of a Green Turtle in Atlantic, 579 
Canada, Pilose Braya, Braya pilosa Hooker (Cruciferae; Bras- 
sicaceae), an Enigmatic Endemic of Arctic, 550 
Canada, Predaceous Water Beetles (Coleoptera: Adephaga: 
Dytiscidae, Gyrinidae) Collected Along the Horton 
and Thelon Rivers in the Arctic Central Barrens of, 
425 
Canada, Sequential Polyandry in Piping Plover, Charadrius 
melodus, Nesting in Eastern, 444 
Canada, Stranding of a Pygmy Sperm Whale, Kogia brevi- 
ceps, in the Northern Gulf of St. Lawrence, 495 
Canada 2004 and 2005, Calendar: Endangered Reptiles of, 
644 
Canadian Arctic, Movements of Subadult Male Grizzly Bears, 
Ursus arctos, in the Central, 239 
Canadian Arctic Islands Using Morphology and Chloroplast 
DNA Data, A Systematic Analysis of the Alpine Saxi- 
frage Complex (Saxifragaceae) in the, 326 
Canadian Field-Naturalist, Advice for Contributors to The, 
158,294,494 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Canadian Rocky Mountains, Predation on Two Mule Deer, 
Odocoileus hemionus, by a Canada Lynx, Lynx 
canadensis, in the Southern, 191 

Canadian Species at Risk, 484,644 

Candelariella sp., 411 

Canis domesticus, 247 

latrans, 41,96,116,243,278,306,341,368,370,606,608 

lupus, 42,57,61,95,115,127,134,194,247,278,350,361, 
368,603,608 

lupus lycaon, 608 

rufus, 608 

Canis latrans, Evidence for the Use of Vocalization to Coor- 
dinate the Killing of a White-Tailed Deer, Odocoileus 
virginianus, by Coyotes, 278 

Canis latrans, in a Heterogeneous Environment, The Influ- 
ence of Prey Availability and Vegetation Character- 
istics on Scent Station Visitation Rates of Coyotes, 
341 

Canis latrans, Severe Chronic Neck Injury Caused by a Snare 
in a Coyote, 243 

Canis lupus, at Katmai National Park and Preserve, Alaska, 
Interactions of Brown Bears, Ursus arctos, and Gray 
Wolves, 247 

Canis lupus, in and near Pukaskwa National Park, Ontario, 
Demographic patterns and limitations of Grey Wolves, 
95 

Canis lupus, in Relation to Forest Harvesting in West-central 
Alberta, Winter Habitat Use by Wolves, 368 

Canis lupus, Pack Density, Record High Wolf, 127 

Canis lupus, South of the St. Lawrence River in Over 100 
Years, First Confirmed Occurrence of a Wolf, 608 

Canis lupus, Unusual Behavior by Bison, Bison bison, 
Toward Elk, Cervus elaphus, and Wolves, 115 

Capelin, 262 

Capreolus capreolus, 134 

Caprimulgus vociferus, 230 

Capsella bursa-pastoris, 560 

Carcinus maenas, 82 

Cardamine bellidifolia, 559 

impatiens, 613 

Carduelis pinus, 378 

tristis, 378 

Caretta caretta, 579 

Caretta caretta, and the First Record of a Green Turtle in At- 
lantic Canada, Hybridization between a Green Turtle, 
Chelonia mydas, and Loggerhead Turtle, 579 

Carex spp., 247,318,400,501 

albo-nigra, 559 

aquatilis, 591 

athrostachya, 169,559 
atrosquama, 559 

capillaris ssp. chlorostachys, 559 
crawfordii, 559 

eleusinoides, 559 

feta, 170 

foenea, 266,559 

glareosa, 266 

hoodii, 266 

houghtonii, 197 

lapponica, 266 

limosa, 559,591 

loliacea, 559 

magellanica ssp. irrigua, 266 
marina, 266 

obtusata, 559 


pachystachya, 266,559 
pellita, 559 

petasata, 559 

praegracilis, 559 

rossii, 559 

stylosa, 558 

supina ssp. spaniocarpa, 559 
sychnocephala, 559 


i} unilateralis, 169 


viridula, 559 
Caribou, 39,192,241,362,602 
Woodland, 57,61,96,119,368 
‘Caribou, Rangifer tarandus caribou, in Saskatchewan, Mor- 
phology of Female Woodland, 119 
: Caripeta divisata, 538 
| Carpiodes cyprinus, 523 
_ Carpodacus purpureus, 378 
Carposinidae, 536 
Carter, B. A Tribute to Thomas Henry Manning (1911-1998), 
618 
Carterocephalus palaemon, 537 
Case, R., 239 
Cassiope tetragona ssp. saximontana, 559 
| Castor canadensis, 40,129,136,248,267,606 
fiber, 434 


| Castor fiber, Collapsing Burrow Causes Death of a Eurasian 


Beaver, 434 
Castors, 136 
Caswell, P., 558 
-Catharus fuscescens, 232 


| Catillaria chalybeia, 408 
| Catocala briseis, 541 


relicta, 541 
semirelicta, 541 
unijuga, 541 
Catocalinae, 541 
Catostomidae, 521 
Catostomus spp., 521 
commersoni, 319,523 


| Cattail, 253,318 


Common, 545,561 


| Cedar, Eastern White, 95,253,361,508 


Yellow, 159 
Centaurea diffusa, 300 
Centipede, 389 
Centrarchidae, 523 
Cephalophus monticola, 185 


| Cepphus grylle, 258 


Cerastium arvense, 559 
| Cerorhinca monocerata, 615 


| Cerorhinca monocerata, Fledglings Fly to the Sea from Their 


Natal Burrows?, Do Rhinoceros Auklet, 615 
| Certhia americana, 384 
Cervus canadensis, 606 

elaphus, 115,132,185,368 

elaphus nelsoni, 132 

nippon, 185 


Cervus elaphus, and Wolves, Canis lupus, Unusual Behavior 


by Bison, Bison bison, Toward Elk, 115 


Cervus elaphus, in Patagonia, Observations of Interactions 


between Puma, Puma concolor, and Introduced Euro- 
pean Red Deer, 132 

Chamaecyparis nootkatensis, 159 

Chamaerhodos erecta ssp. nuttallii, 559 


| Chapdelaine, G., 256 


INDEX TO VOLUME 118 


655 


Chapleau, F., 521 
Charadrius alexandrinus, 445 
melodus, 444,448 
vociferus, 445 
Charadrius melodus, egg viability after seawater immersion, 
Piping Plover, 448 
Charadrius melodus, Nesting in Eastern Canada, Sequential 
Polyandry in Piping Plover, 444 
Charlock, 572 
Chat-fou liséré, 523 
Chelonia mydas, 72,579,611 
Chelonia mydas, and Loggerhead Turtle, Caretta caretta, and 
the First Record of a Green Turtle in Atlantic Canada, 
Hybridization between a Green Turtle, 579 
Chelydra serpentina, 610 
Chelydra serpentina?, Is Cost of Locomotion the Reason for 
Prolonged Nesting Forays of Snapping Turtles, 610 
Chenopodium capitatum, 559 
Cherry, Choke, 253,362 
Sand, 196 
Chevalier blanc, 523 
de riviere, 521 
doré 526 
rouge, 523 
Chickadee, Black-capped, 232 
Boreal, 48 
Chestnut-backed, 378 
Chickweed, Field, 569 
Chilocorus stigma, 312 
Chionodes continuella, 534 
lugubrella, 534 
mediofuscella, 534 
occlusus, 534 
psilopterus, 534 
sattleri, 534 
terminimaculella, 534 
Chipmunk, Yellow-pine, 299 
Chipmunks, Tamias amoenus, in Old Field and Orchard Habi- 
tats, Population Dynamics of Deer Mice, Peromyscus 
maniculatus, and Yellow-pine, 299 
Chondrus crispus, 226 
Chorda filum, 226 
Chordaria flagelliformis, 226 
Choreutidae, 536 
Choreutis diana, 536 
Choristoneura albaniana, 535 
conflictana, 535 
fumiferana, 361,535 
rosaceana, 535 
Choristostigma plumbosignale, 537 
Chrysanympha formosa, 541 
Chytolita petrealis, 540 
Chytonix palliatricula, 542 
Cicuta maculata vat. angustifolia, 559 
Cinclus mexicanus, 443,446 
Cinclus mexicanus, Preys Upon Larval Tailed Frogs, Asca- 
phus truei, American Dipper, 446 
Cinquefoil, Tall, 573 
Ciona intestinalis, 79 
Ciprés, 132 
Cirsium muticum, 253 
Cirtwill, A., 604 
Cladara atroliturata, 
limitaria, 539 
Cladina rangiferina, 


539 


591 


656 


Clam, Butter, 77 
Manila, 78 
Native Littleneck, 77 
Soft-shell, 78 
Clangula hyemalis, 604 
Clangula hyemalis, Two Great Black-backed Gulls, Larus 
marinus, Kill Male Longtailed Duck, 604 
Clayburn, J.K., 615 
Claytonia megarrhiza, 558 
Cleavers, 175 
Clemensia albata, 540 
Clepsis clemensiana, 535 
melaleucana, 535 
persicana, 535 
virescana, 535 
Clintonia borealis, 362 
Clintonia, Yellow, 362 
Cliostomum griffithii, 424 
Clostera albosigma, 540 
apicalis, 540 
strigosa, 540 
Clover, 253,362 
Red, 574 
Cluff, H.D., 239 
Coad, B.W., Reviews by, 281,282,469 
Cobb, L.M. and V.A. Cobb. Occurrence of Parasitoid Wasps, 
Baeus sp. and Gelis sp., in the Egg Sacs of the Wolf 
Spiders Pardosa moesta and Pardosa sternalis (Ara- 
neae, Lycosidae) in Southeastern Idaho, 122 
Cobb, V.A., 122 
Coccinella septempunctata, 225,309 
transversoguttata, 310 
transversoguttata richardsonii, 309 
Coccotrema cucurbitula, 420 
maritimum, 406 
Cochylis nana, 535 
Cody, W.J., C.E. Kennedy, B. Bennett, and P. Caswell. New 
Records of Vascular Plants in the Yukon Territory VI, 
558 
Cody, W.J., Reviews by, 147,289,476 
Coenophila opacifrons, 543 
Coir6n Amargo, 132 
Dulce, 132 
Colaptes auratus, 358 
Coleophora sp., 533 
alnifoliae, 533 
comptoniella, 533 
corylifoliella, 533 
dextrella, 533 
duplicis, 533 
glaucicolella, 533 
mayrella, 533 
mcedunnoughiella, 533 
persimplexella, 533 
pruniella, 533 
rosaevorella, 533 
Coleophoridae, 533 
Coleotechnites atrupictella, 534 
blastivora, 534 
florae, 534 
piceaella, 534 
Collema fecundum, 411 
novozelandicum, 420 
Colletia spinosissima, 132 
Comptonia peregrina, 196 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Comtois, A., F. Chapleau, C.B. Renaud, H. Fournier, B. Camp-) 
bell, et R. Pariseau. Inventaire printanier d’une frayére 
multispécifique: l’ichtyofaune des rapides de la riv- 
iére Gatineau, Québec, 521 

Contopus cooperi, 378 

virens, 232 

Cook, E.R., Reviews by, 138,141,467 

Cook, J.A., 438 

Copepod, 77 

Corispermum hyssopifolium sensu, 568 

ochotense var. alaskanum, 558 
Cormorant, Double-crested, 257 
Great, 257 

Corn, 253 

Cornus canadensis, 46 
racemosa, 342 
stolonifera, 253 

Coronopus didymus, 613 

Corophium acherusicum, 77 

Corvus brachyrhynchos, 232 

caurinus, 378 
corax, 273 

Corylus cornuta, 197,251 

COSEWIC Assessment Results November 2003, 155 

Cosmopterigidae, 534 

Cossidae, 534 

Cotton-grass, Short-anthered, 566 

Cottontail, Eastern, 342 

Cottonwood, Black, 211,355 

Plains, 309 

Cottus bairdi, 323 

cognatus, 323 

Cotula coronopifolia, 169 

Couette, 523 

Cougar, 159,370 

Cougars, Puma concolor vancouverensis, Morphology and 
Population Characteristics of Vancouver Island, 159 

Couleuvre brune, 136 

mince, 135 
rayée de l’est, 135 

Couleuvre mince, Thamnophis sauritus septentrionalis, au 
Québec, Premiéres mentions de la, 135 

Cowbird, Brown-headed, 378 

Coyote, 41,96,116,243,278,306,341,368,606,608 

Western, 608 

Coyote, Canis latrans, Severe Chronic Neck Injury Caused by 
a Snare in a, 243 

Coyotes, Canis latrans, Evidence for the Use of Vocalization 
to Coordinate the Killing of a White-Tailed Deer, 
Odocoileus virginianus, by, 278 

Coyotes, Canis latrans, in a Heterogeneous Environment, 
The Influence of Prey Availability and Vegetation 
Characteristics on Scent Station Visitation Rates of, 
341 

Crab, European Green, 82 

Crambidae, 536 

Crambus leachellus, 536 

perlellus, 536 

Crapet arlequin, 523 

de roche, 523 
soleil, 523 

Crassostrea gigas, 77 

Crataegus sp., 365 

Creeper, Brown, 378 

Cress, Penny, 573 


ii) 


| 


2004 


| Créte, M., 360 
| Crow, American, 232 


Northwestern, 378 


| Cryptocala acadiensis, 543 
| Cryptococcus spp., 41 


Cryptogramma crispa vat. acrostichoides, 559 
Cryptosula pallasiana, 79 

Cuculliinae, 541 

Cudweed, Lowland, 169 


/ Cumming, E.E. Habitat Segregation Among Songbirds in 


Old-Growth Boreal Mixedwood Forest, 45 
Cyanocitta cristata, 231 
Cyclophora pendulinaria, 538 


Cydia flexiloqua, 536 


populana, 536 


| Cynosurus echinatus, 176 


Cyperus sp., 180 
squarrosus, 179 
Cyperus, Awned, 179 


| Cyprinidae, 523 


Cytisus scoparius, 177 


Dactylis glomerata, 174,300 

‘Daigle, C., M. Créte, L. Lesage, J.-P. Ouellet, and J. Huot. 
Summer Diet of Two White-tailed Deer, Odocoileus 
virginianus, Populations Living at Low and High Den- 
sity in Southern Québec, 360 


| Danthonia californica, 169 


spicata, 277 


_| Daoust, P.-Y. and P.H. Nicholson. Severe Chronic Neck Injury 


Caused by a Snare in a Coyote, Canis latrans, 243 


| Darbyshire, S.J. The Northern True Katydid, Pterophylla 


camellifolia (Orthoptera: Pseudophyllidae), at Ottawa, 
Ontario, 124 


) Darter, Iowa, 319 


Johnny, 319 
Rainbow, 319 


| Dasychira plagiata, 540 


vagans, 540 
Daucus carota, 342 
Deer, 368 
Columbia Black-tailed, 162 
European Red, 132 
Mule, 132,162,185,191,370,606 
Red, 185 
Roe, 134 
Sika, 185 
White-tailed, 40,96, 127,185,278,360,370,606 


| Deer, Cervus elaphus, in Patagonia, Observations of Inter- 


actions between Puma, Puma concolor, and Introduced 
European Red, 132 

Odocoileus hemionus, by a Canada Lynx, Lynx cana- 
densis, in the Southern Canadian Rocky Mountains, 

| Predation on Two Mule, 191 

Deer, Odocoileus virginianus, by Coyotes, Canis latrans, 


| Deer, 


Evidence for the Use of Vocalization to Coordinate 
the Killing of a White-Tailed, 278 

Odocoileus virginianus, Populations Living at Low 
and High Density in Southern Québec, Summer Diet 
of Two White-tailed, 360 


| Deer, 


| Deer, Odocoileus virginianus, with Medetomidine and Ket- 


amine, and Antagonism with Atipamezole, Immobi- 
lization of Clover-trapped White-tailed, 185 


| Dendroica castanea, 48 


cerulea, 229 


INDEX TO VOLUME 118 


657 


coronata, 48,86,232,378 
fusca, 48 
magnolia, 48 
pensylvanica, 232 
petechia, 86,232 
virens, 48,232 
Dendroica cerulea Management on Eastern Ontario Bird 
Species, Predicting the Effects of Cerulean Warbler, 
229 
Depressariodes ciniflonella, 533 
Dermacentor variabilis, 42 
Dermochelys coriacea, 72,579 
Deschampsia brevifolia, 559 
Descurainia incisa vat. incisa, 559 
Desmognathus ochrophaeus, 136 
de Solla, S.R. and K.J. Fernie. Is Cost of Locomotion the Rea- 
son for Prolonged Nesting Forays of Snapping Turtles, 
Chelydra serpentina?, 610 
Desroches, J.-F. et I. Picard. Extension de I’ aire de distribution 
connue de la Musaraigne fuligineuse, Sorex fumeus, 
dans le nord-est du Québec, 441 
Desroches, J.-F. et R. Lapar. Premiéres mentions de la Cou- 
leuvre mince, Thamnophis sauritus septentrionalis, 
au Québec, 135 
Diachrysia aereoides, 541 
Diarsia dislocata, 542 
rosaria, 542 
rubifera, 542 
Dibb, A., 191 
Dicamptodon copei, 448 
ensatus, 448 
Dichomeris levisella, 534 
Dicrostonyx hudsonius, 403 
richardsoni, 403 
Digrammia rippertaria, 537 
Dioryctria reniculelloides, 536 
Dipper, American, 443,446 
Dipper, Cinclus mexicanus, Preys Upon Larval Tailed Frogs, 
Ascaphus truei, American, 446 
Dock, Curled, 568 
Dodecatheon hendersonii, 175 
hendersonii ssp. hendersonii, 175 
Dogtail, Hedgehog, 176 
Dogwood, 253 
Gray, 342 
Dolichomia thymetusalis, 536 
Doré jaune, 523 
noir, 523 
Dorosoma cepedianum, 318 
Douglas, G.W., 179 
Douglas, G.W. Conservation Evaluation of Lemmon’s Holly 
Fern, Polystichum lemmonii, a Threatened Fern in 
Canada, 164 
Douglas, G.W. and J.L. Penny. Conservation Evaluation of 
Howell’s Triteleia, Triteleia howellii, an Endangered 
Lily in Canada, 174 
Douglas, G.W. and J.M. Illingworth. Conservation Evalua- 
tion of the Pacific Population of Tall Woolly-heads, 
Psilocarphus elatior, an Endangered Herb in Canada, 
169 
Douglasia alaskana, 559 
arctica, 558 
Dowsley, M. and A. Cirtwill. Two Great Black-backed Gulls, 
Larus marinus, Kill Male Longtailed Duck, Clangu- 
la hyemalis, 604 


658 


Draba aurea, 559 
corymbosa, 550,559 
crassifolia, 559 
glabella, 559 
nemorosa, 559 
ogilviensis, 559 
palanderiana, 559 
porsildii, 559 
stenoloba, 559 
Draba, Golden, 571 
Drepana arcuata, 537 
bilineata, 537 
Drepanidae, 537 
Dryas hookeriana, 559 
Duchesne, L.C. and S. Wetzel. Effect of Fire Intensity and 
Depth of Burn on Lowbush Blueberry, Vaccinium 
angustifolium, and Velvet Leaf Blueberry, Vaccinium 
myrtilloides, Production in Eastern Ontario, 195 
Duck, American Black, 129,604 
Black, 268 
Longtailed, 604 
Ring-necked, 268 
Ruddy, 604 
Wood, 131,230 
Duck, Anas rubripes, Broods, Extraordinary Size and Survival 
of American Black, 129 
Duck, Clangula hyemalis, Two Great Black-backed Gulls, 
Larus marinus, Kill Male Longtailed, 604 
Ducks, Barrow’s Goldeneye, Bucephala islandica, and Surf 
Scoter, Melanitta perspicillata, in Coastal South- 
western British Columbia, Observations of Above- 
Surface Littoral Foraging in Two Sea, 264 
Duiker, Blue, 185 
Dulse, 226 
Dutton, P.H., 579 
Dyke, A.S., D.A. Hodgson, E.L. Bousfield, and R.E. Bedford. 
A Tribute to Victor Kent Prest (1913-2003), 626 
Dysstroma citrata, 538 
formosa, 538 
hersiliata, 538 
truncata, 538 
walkerata, 538 


Eagle, Bald, 267 
Earthworm, 388 
Ecliptopera silaceata, 538 
Ectropis crepuscularia, 537 
Editor’s Report for Volume 117 (2003), 296 
Eelgrass, 225,318 
Eelgrass, Zostera marina, Wrack by Three Species of Lady- 
bird Beetles (Coleoptera: Coccinellidae) in Prince 
Edward Island, Use of, 225 
Egira dolosa, 542 
Eider, Common, 258 
Eilema bicolor, 540 
Elachista sp., 533 
adempta, 533 
albicapitella, 533 
Elachistidae, 533 
Elaphria alapallida, 542 
Elderberry, Red, 577 
Eleocharis elliptica, 266 
uniglumis, 559 
Elk, 115,368 
American, 132,606 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Elk, Cervus elaphus, and Wolves, Canis lupus, Unusual 
Behavior by Bison, Bison bison, Toward, 115 
Elm, American, 310,508 
Elodea canadensis, 318 
Elymus alaskanus ssp. alaskanus, 559 
glaucus, 559 
macrourus, 559 
repens, 253 
spicatus, 559 
trachycaulus ssp. andinus, 559 
Elytrigia intermedia, 560 
Empidonax difficilis, 378 
minimus, 86,232 
Emydura macquarii, 610 
Enargia decolor, 542 
infumata, 542 
Engeman, R.M., 201 
Enhydra lutris, 270 
Enhydra lutris, Sightings off Haida Gwaii/Queen Charlotte 
Islands, British Columbia, 1972-2002, Sea Otter, 270 © 
Ennominae, 537 
Ennomos magnaria, 538 
Eosphoropteryx thyatyroides, 541 
Ephebe lanata, 424 
Epilobium angustifolium, 253,365 
Epinotia castaneana, 536 
criddleana, 536 
lindana, 536 
momonana, 536 
nisella, 536 
rectiplicana, 536 
solandriana, 536 
transmissana, 536 
trigonella, 536 
Epirrhoe alternata, 539 
Epirrita autumnata, 539 
Epp, B. and J.C. Tardif. Effects of Lodgepole Pine Dwarf 
Mistletoe, Arceuthobium americanum, on Jack Pine, 
Pinus banksiana, Growth in Manitoba, 595 
Equisetum spp., 46 
variegatum, 591 
Equus cabalus, 370 
Erannis tiliaria, 538 
Erethizon dorsatum, 607 
Eretmochelys imbricata, 580 
Erickson, W.R. Bird Communities of the Garry Oak Habitat 
in Southwestern British Columbia, 376 
Erigeron caespitosus, 559 
hyperboreus, 559 
Eriophorum brachyantherum, 559 
Eritrichium aretioides, 559 
chamissonis, 559 
Ermine, 243 
Ernst, J., 56,61 
Errata for The Canadian Field-Naturalist 118(3), 649 
Erratum Canadian Field-Naturalist 118(2), 485 
Esocidae, 523 
Esox lucius, 523 
masquinongy, 523 
Espino Negro, 132 
Esturgeon jaune, 523 
Etheostoma caeruleum, 319 
exile, 319 
nigrum, 319,523 
olmstedi, 523 


2004 


| 
Eubaphe mendica, 539 
Euchlaena marginaria, 538 
obtusaria, 538 

| tigrinaria, 538 
_Eudonia albertalis, 536 
| Eueretagrotis perattenta, 543 
_Euhyponomeutoides gracilariella, 532 
Eulia ministrana, 535 
| Eulithis explanata, 538 
propulsata, 538 
| xylina, 538 

Eulogia ochrifrontella, 536 
_Euonymus fortunei, 614 
| Euphrasia subarctica, 559 
| Euphyia intermediata, 539 
| Eupithecia anticaria, 539 
i| assimilata, 539 

columbiata, 539 
i| perfusca, 539 
ravocostaliata, 539 
satyrata, 539 
stellata, 539 

| subfuscata, 539 
| Euplexia benesimilis, 542 
| Eurois astricta, 543 
i occulta, 543 
 Eustrotiinae, 541 
Euthyatira pudens, 537 

| Everes amyntula, 537 

Everett, K.T. and K.A. Otter. Presence of Cavities in Snags 

Retained in Forest Cutblocks: Do Management Poli- 

i cies Promote Species Retention?, 354 
| | Evergestis pallidata, 537 


Fagus, 52 

i grandifolia, 41,361,507 

Falseflax, 571 

| Feathermoss, Red-stemmed, 591 

| Felis concolor, 370 

| Ferguson, C., 225 

Ferguson, S.H. and D.K.A. Berube. Invertebrate Diversity 

under Artificial Cover in Relation to Boreal Forest 
Habitat Characteristics, 386 

“Fern, 159 

| Bracken, 196,251 

Kruckeberg’s Holly, 164 

| Lemmon’s Holly, 164 

Mountain Holly, 164 

| Nahanni Oak, 560 

| Ostrich, 46,253 


| Sensitive, 253 

Sweet, 196 

| Fern, Polystichum lemmonii, a Threatened Fern in Canada, 
Conservation Evaluation of Lemmon’s Holly, 164 
| Fernie, K.J., 610 

| Fescue, Rocky Mountain, 563 

i Short-leaf, 562 

. Festuca brachyphylla, 559 

pallescens, 132 

| richardsonii, 559 

saximontana, 559 

Filatima abactella, 534 

_ Finch, Purple, 378 

Fir, Balsam, 46,95,251,268,274,361,388,401,514,530 
Douglas, 159,355,376,584,595 


| 
i 
| 


INDEX TO VOLUME 118 


659 


Sub-alpine, 191,355,584 
Fireweed, 253 
Fisher, 56,96,583 
Flicker, Common, 379 
Northern, 358,378 
Flounder, Starry, 248 
Flueck, W.T. Observations of Interactions between Puma, 
Puma concolor, and Introduced European Red Deer, 
Cervus elaphus, in Patagonia, 132 
Flycatcher, Least, 232 
Olive-sided, 378 
Pacific-coast, 378 
Pacific-slope, 378 
Least, 86 
Fool’s-parsley, 613 
Forget-me-not, 576 
Forshner, S.A., P.C. Paquet, F.G.M. Burrows, G.K. Neale, 
K.D. Wade and W.M. Samuel. Demographic patterns 
and limitations of Grey Wolves, Canis lupus, in and 
near Pukaskwa National Park, Ontario, 95 
Fouille-roche gris, 523 
zébré, 523 
Fournier, H., 521 
Fox, 58,273 
Arctic, 602 
Red, 40,69,96,244,249 368 
Fox, Alopex lagopus, den by a Polar Bear, Ursus maritimus, 
Excavation of an Arctic, 602 
Fragaria sp., 365 
Fraser, S., 225 
Fratercula arctica, 258,604,615 
cirrhata, 615 
corniculata, 615 
Fraxinus sp., 136,253 
americana, 361,508 
nigra, 310,365 
pennsylvanica, 309 
Frog, Red-legged, 448 
Tailed, 447 
Froglog: Newsletter of the Declining Amphibian Populations 
Task Force, 155,294,484,645 
Frogs, Ascaphus truei, American Dipper, Cinclus mexicanus, 
Preys Upon Larval Tailed, 446 
Fucus gardneri, 406 
serratus, 226 
Fungus, Nectria, 41 
Furcellaria lumbricalis, 226 
Furcula modesta, 540 
occidentalis, 540 
scolopendrina, 540 
Fuscidea cyathoides, 420 
intercincta, 421 
mollis, 424 
thomsonii, 406 
Fuscopannaria leucostictoides, 411 
maritima, 411 
thiersii, 421 


Galeopsis tetrahit ssp. bifida, 560 

Galium aparine, 175 

Garbary, D.J., S. Fraser, C. Ferguson, and R.F. Lauff. Use of 
Eelgrass, Zostera marina, Wrack by Three Species 
of Ladybird Beetles (Coleoptera: Coccinellidae) in 
Prince Edward Island, 225 


660 


Gau, R.J., B.D. McLoughlin, R. Case, H.D. Cluff, R. Mulders, 
and F. Messier. Movements of Subadult Male Grizzly 
Bears, Ursus arctos, in the Central Canadian Arctic, 
239 
Gaultheria procumbens, 196 
shallon, 159 
Gavia spp., 210 
immer, 210 
pacifica, 210 
stellata, 258 
Gavia spp.) and Red-necked Grebe (Podiceps grisegena) 
Populations in the Lower Matanuska-Susitna Valley, 
Alaska, Changes in Loon, 210 
Gawn, M., Review by, 473 
Gelechia dyarella, 534 
lynceella, 534 
Gelechiidae, 534 
Gelis sp., 122 
Gelis sp., in the Egg Sacs of the Wolf Spiders Pardosa moesta 
and Pardosa sternalis (Araneae, Lycosidae) in South- 
eastern Idaho, Occurrence of Parasitoid Wasps, Baeus 
sp. and, 122 
Gemsbok, 185 
Gende, S.M., 499 
Gentianella tenella, 559 
Geometridae, 537 
Geothlypis trichas, 86,232 
Geum aleppicum ssp. strictum, 559 
rossti, 559 
Giberson, D.J., 425 
Gillespie, L.J., 326 
Ginns, J., Review, by, 146 
Gladders, A., 159 
Glaux maritimus, 226 
Glen, W.M., 504 
Gluphisia avimacula, 540 
lintneri, 540 
septentrionis, 540 
Glyceria striata var. stricta, 559 
Glycine max, 253 
Gnaphalium palustre, 169 
Gnatcatcher, Blue-grey, 232 
Gnorimoschema septentrionella, 534 
vastificum, 534 
Goat, Mountain, 370 
Goby, Round, 318 
Tubenose, 318 
Goby, Neogobius melanostomus Pallas (Gobiidae), and Eco- 
tone Utilisation in St. Clair Lowland Waters, Ontario, 
Description of Age-O0 Round, 318 
Goh, K.M.L., 159 
Goldeneye, 604 
Barrow’s, 264 
Common, 267 
Goldeneye, Bucephala islandica, and Surf Scoter, Melanitta 
perspicillata, in Coastal Southwestern British Colum- 
bia, Observations of Above-Surface Littoral Foraging 
in Two Sea Ducks, Barrow’s, 264 
Goldenrod, 253 
Tall, 342 
Goldfinch, American, 378 
Goodwin, C.E., Review by, 138 
Goose, Canada, 268,602 
Snow, 602 
Gracillariidae, 532 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Grand brochet, 523 
Graphiphora augur, 543 
Grapholita lunatana, 536 


Grass, 400 
Annual Blue, 563 
Cheat, 174 


Creeping Bent, 562 
Crested Wheat, 561 
Forked Three-awned, 276 
Fowl Marina, 563 
Indian, 342 
June, 563 
Orchard, 174,300 
Quack, 300 
Western Rye, 562 
Wheat, 584 
Grass, Aristida basiramea Engelm. ex Vasey: A New Addition 
to the Flora of Quebec, Forked Three-awned, 276 
Grayling, Arctic, 111 
Grayling, Thymallus arcticus, Consumption of Shrews, Sorex 
spp., by Arctic, 111 
Grebe, Red-necked, 210 
Grebe (Podiceps grisegena) Populations in the Lower Mata- 
nuska-Susitna Valley, Alaska, Changes in Loon (Gavia 
spp.) and Red-necked, 210 
Greya politella, 532 
Griselda radicana, 536 
Grosbeak, Rose-breasted, 232 
Grouse, 41 
Ruffed, 232 
Guillemot, Black, 257 
Gull, 249,442 
Bonaparte’s, 443 
Glaucous-winged, 443,616 
Great Black-backed, 257,604 
Herring, 257,443,449 
Mew, 443 
Ring-billed, 258 
Gulls, Larus marinus, Kill Male Longtailed Duck, Clangula 
hyemalis, Two Great Black-backed, 604 
Gulls, Larus spp., Foraging at Pink Salmon, Oncorhynchus 
gorbuscha, Spawning Runs, 442 
Gulo gulo luscus, 56,61 
Gulo gulo luscus, in the boreal forest, Effects of mid-winter 
snow depth on stand selection by Wolverines, 56 
Gulo gulo luscus, Resting Sites and Caching Behavior in the 
Boreal Forest, Wolverine, 61 
Gyalecta jenensis, 406 
Gymnocarpium jessoense ssp. parvulum, 559 
Gypsonoma adjuncta, 536 
fasciolana, 535 
salicicolana, 536 
substitutionis, 536 
Gyrinus opacus, 431 


Habrosyne scripta, 537 

Hadeninae, 541 

Haemoproteus, 236 

Hahn, A., 159 

Hairgrass, Early, 177 
Mountain, 564 

Haliaeetus leucocephalus, 267 

Hamr, J., 251 


2004 


Hansen, D.J. Observations of Habitat Use by Polar Bears, 
Ursus maritimus, in the Alaskan Beaufort, Chukchi, 
and Northern Bering Seas, 395 

Hansen, L.P., 185 

Haploa lecontei, 540 

Hare, 607 

Snowshoe, 41,58,191,278,585 

Hare, J.F., G. Todd, and W.A. Untereiner. Multiple Mating 
Results in Multiple Paternity in Richardson’s Ground 
Squirrels, Spermophilus richardsonii, 90 


| Harrimanella stellariana, 559 


Harris, A.G., 179 

Harris, J.G. Pilose Braya, Braya pilosa Hooker (Cruciferae; 
Brassicaceae), an Enigmatic Endemic of Arctic Cana- 
da, 550 

Harris, S.A. and P.W. Ball. New Records of Cyperaceae and 
Juncaceae from the Yukon Territory, 266 

Harrisimemna trisignata, 541 

Hawk, Rough-legged, 268 


_ Hay, 361 
| Hayward, J.L. and J.K. Clayburn. Do Rhinoceros Auklet, 


Cerorhinca monocerata, Fledglings Fly to the Sea 
from Their Natal Burrows?, 615 
Hazel, Beaked, 251 


_ Healy, C. and L.J. Gillespie. A Systematic Analysis of the 


Alpine Saxifrage Complex (Saxifragaceae) in the Can- 
adian Arctic Islands Using Morphology and Chloro- 
plast DNA Data, 326 

Hedeoma hispida, 277 

Helcystogramma fernaldella, 534 

Hellinsia homodactylus, 536 


_ Hemicarpha, 179 
| Hemlock, 41,361,499 


Eastern, 251,508 
Mountain, 159 
Western, 159 
Hemp-nettle, 576 
Hepialidae, 532 
Heracleum, 501 
Herminiinae, 540 
Heron, Great Blue, 230 
Herteliana alaskensis, 408 
Hesperiidae, 537 
Heterocodon rariflorum, 170 
Heterocodon, 170 
Hillia iris, 542 
Hinam, H.L., S.G. Sealy, and T.J. Underwood. Ruby-throated 
Hummingbird, Archilochus colubris, Entanglements 
in Burdock, Arctium spp., at Delta Marsh, Manitoba, 
85 


| Hiodon tergisus, 523 


Hiodontidae, 523 
Hippodamia convergens, 309 
parenthesis, 309 
tredecimpunctata, 225,309 
Hippotragus equinus, 185 
Hirundo rustica, 230 
Hodgson, D.A., 626 
Hogue, E.J., 299 
Holodiscus discolor, 176 
Homan, H.J., G.M. Linz, R.M. Engeman, and L.B. Penry. 
Spring Dispersal Patterns of Red-winged Blackbirds, 
Agelaius phoeniceus, Staging in Eastern South Dako- 
ta, 201 
Homarus americanus, 82 


INDEX TO VOLUME 118 


66] 


Homoglaea hircina, 542 
Homohadena badistriga, 5: 
infixa, 541 
Homosetia fasciella, 532 
Honeysuckle, Hairy, 176 
Hood, G.A. and T. Neufeld. First Record of Mountain Lions, 
Puma concolor, in Elk Island National Park, Alberta, 
605 
Hop-clover, Small, 170 
Hordeum brachyantherum, 559 
vulgare, 361 
Horse, Wild, 370 
Houston, C.S., Reviews by, 142,290,465,466,470,47 1,481,631, 
633,635 
Huckleberry, Blue, 198 
Huettmann, F., Reviews by, 478,632 
Hummingbird, Ruby-throated, 85 
Rufous, 378 
Hummingbird, Archilochus colubris, Entanglements in Bur- 
dock, Arctium spp., at Delta Marsh, Manitoba, Ruby- 
throated, 85 
Huot, J., 360 
Hydrelia albifera, 539 
Hydriomena furcata, 539 
perfracta, 538 
renunciata, 539 
ruberata, 539 
Hydroporus appalachius, 431 
geniculatus, 431 
hudsonicus, 432 
morio, 431 
Hydrozoan, 78 
Hygrotus novemlineatus, 430 
HAyles gallii, 540 
Hylocichla mustelina, 232 
Hylocomium splendens, 591 
Hypatopa titanella, 533 
Hypena atomaria, 541 
edictalis, 541 
humuli, 541 
Hypeninae, 541 
Hypenodes fractilinea, 540 
Hyperaspis benedetti, 312 
Hyppa contrasta, 542 


Ichthyomyzon fossor, 523 
unicuspis, 523 

Ictaluridae, 523 

Ictalurus punctatus, 523 

Icterus galbula, 232 

Idaea rotundopennata, 538 

Idaho, Occurrence of Parasitoid Wasps, Baeus sp. and Gelis 
sp., in the Egg Sacs of the Wolf Spiders Pardosa moesta 
and Pardosa sternalis (Araneae, Lycosidae) in South- 
eastern, 122 

Idia aemula, 540 

americalis, 540 

Illex illecebrosus, 497 

Illingworth, J.M., 169 

Ilybius erichsoni, 431 

Impala, 185 

Indian’s Dream, 166 

Ipimorpha pleonectusa, 542 

Tridopsis larvaria, 537 

Tris setosa ssp. interior, 559 


662 


Iris, Wild, 567 

Isoetes ?maritima, 559 
echinospora, 559 

Isopod, 389 


James, M.C., K. Martin, and P.H. Dutton. Hybridization be- 
tween a Green Turtle, Chelonia mydas, and Logger- 
head Turtle, Caretta caretta, and the First Record of 
a Green Turtle in Atlantic Canada, 579 

Jay, Blue, 231 

Gray, 273 

John, R., Reviews by, 140,143,147,150,282,288,291,630,631, 
634,636 

Joint, Blue, 591 

Jolicoeur, H., 608 

Jones, J., W.J. McLeish, and R.J. Robertson. Predicting the 
Effects of Cerulean Warbler, Dendroica cerulea Man- 
agement on Eastern Ontario Bird Species, 229 

Junco hyemalis, 235,381 

hyemalis cismontanus, 235 
hyemalis hyemalis, 235 
hyemalis oregonus, 235 

Junco, Dark-eyed, 235,381 

Junco hyemalis, Breeding at Different Elevations, Parasite 
Prevalence in Dark-eyed Juncos, 235 

Juncos, Junco hyemalis, Breeding at Different Elevations, 
Parasite Prevalence in Dark-eyed, 235 

Juncus spp., 400 

bufonius, 170,559 
filiformis, 559 
Stygius SSp. americanus, 267 
Juniper, Common, 166 
Juniperus communis, 166 


Kalmia, 501 
angustifolia, 196 
Katydid, Northern True, 124 
Katydid, Prerophylla camellifolia (Orthoptera: PseudophyI- 
lidae), at Ottawa, Ontario, The Northern True, 124 
Kemp, J.P. and C.R. Lacroix. Estimation of Seed Bank and 
Seed Viability of the Gulf of Saint Lawrence Aster, 
Symphyotrichum laurentianum, (Fernald) Nesom, 105 
Kenagy, G.J., 111 
Kennedy, C.E., 558 
Kent, D., 264 
Kierstead, J., 444 
Killdeer, 445 
Kinglet, Golden-crowned, 48,86 
Ruby-crowned, 48,86 
Kittiwake, Black-legged, 257 
Knapweed, Diffuse, 300 
Kneteman, J., 368 
Kobresia simpliciuscula, 559 
Kobresia, Simple, 566 
Koeleria macrantha, 559 
Kogia breviceps, 495 
simus, 495 
Kogia breviceps, in the Northern Gulf of St. Lawrence, Cana- 
da, Stranding of a Pygmy Sperm Whale, 495 
Kohlmeyera complicatula, 420,424 
Korscheltellus gracilis, 532 
Kuzyk, G.W., J. Kneteman, and F.K.A. Schmiegelow. Winter 
Habitat Use by Wolves, Canis lupus, in Relation to 
Forest Harvesting in West-central Alberta, 368 


THE CANADIAN FIELD-NATURALIST 


l’Esturgeon jaune, 521 
Labidesthes sicculus, 319 
Labrador, “Ashkui” Vernal Ice-cover Phenomena and Their | 
Ecological Role in Southern, 267 
Labrador, First Record of a Barred Owl, Strix varia, in, 273 
Lacanobia radix, 542 
Lace, Queen Anne’s, 342 
Lacinipolia lorea, 542 
renigera, 542 
Lacroix, C.R., 105 
Lacroix, D.L., K.G. Wright, and D. Kent. Observations of ' 
Above-Surface Littoral Foraging in Two Sea Ducks, . 
Barrow’s Goldeneye, Bucephala islandica, and Surf | 
Scoter, Melanitta perspicillata, in Coastal Southwest- 
ern British Columbia, 264 
Lactuca serriola, 300 
Ladies’-tresses, Hooded, 567 
Lady’s-fingers, 613 
Lagopus lagopus, 602 
Lambdina fiscellaria, 538 
Lamna nasus, 245 
Lampetra appendix, 523 
Lamproie argentée, 523 
de lest, 523 
du nord, 523 
Lamy, R., 270 
Landry, J.-F., 530 
Langor, D.W., 530 
Lapar, R., 135 
Laquaiche argentée, 523 
Larch, Western, 595 
Larentiinae, 538 
Lariviere, S., Review by, 139 
Larix laricina, 63,136,370,510,590 
Larus spp., 249,442 
argentatus, 258,443,449 
canus, 443 
delawarensis, 258 
glaucescens, 443,616 
marinus, 258,604 
philadelphia, 443 
Larus marinus, Kill Male Longtailed Duck, Clangula hye- 
malis, Two Great Black-backed Gulls, 604 
Larus spp., Foraging at Pink Salmon, Oncorhynchus gor- 
buscha, Spawning Runs, Gulls, 442 
Lasiocampidae, 539 
Lasionycta poca, 542 
antigReke225 
Lauff, R.F., Reviews by, 144,145,288,630 
Laura, 132 
Laurel, Sheep, 196 
Lecanora sp., 411 
actophila, 420 
contractula, 424 
dispersa, 421 
helicopsis, 420 
muralis, 411 
poliophaea, 424 | 
poliophaea ssp. poliophaea, 421 
straminea, 411 
tenera, 424 
. 
: 


Lecidea conferenda, 424 
Lecidella scabra, 424 

stigmatea, 411 

stigmatea f. stigmatea, 424 


Mi 


— 


2004 


Ledum groenlandicum, 591 

Lemming, Brown, 440 
Varying, 403 

Lemmus trimucronatus, 440 

Lenga, 132 


| Lepidium densiflorum var. densiflorum, 559 


densiflorum var. macrocarpum, 560 
Lepidochelys kempii, 580 
Lepomis gibbosus, 523 
macrochirus, 318,523 
Leptinotarsa decemlineata, 226 
Lepus spp., 607 
americanus, 41,58,191,278,585 
Lesage, L., 360 
Leslie, J.K. and C.A. Timmins. Description of Age-O Round 
Goby, Neogobius melanostomus Pallas (Gobiidae), 
and Ecotone Utilisation in St. Clair Lowland Waters, 
Ontario, 318 


| Lettuce, Prickly, 300 


Leucania insueta, 542 
Leucocytozoon, 236 
Lichen, 408 

Grey Reindeer, 591 
Lichina pygmaea, 419 
Lily-of-the-Valley, Wild, 362,567 


Lim, B.K., Reviews by, 286,472 


Limnaecia phragmitella, 534 


_ Limnodrilus monothecus, 78 


Limonium nashii, 226 

Lingonberry, 591 

Linz, G.M., 201 

Lion, Mountain, 159,605 

Lions, Puma concolor, in Elk Island National Park, Alberta, 
First Record of Mountain, 605 

Lipocarpha micrantha, 179 

Lipocarpha, Small-flowered, 179 

Lipocarpha, Lipocarpha micrantha (Cyperaceae), in Canada, 
Conservation Evaluation of Small-flowered, 179 

Lipocarpha micrantha (Cyperaceae), in Canada, Conservation 
Evaluation of Small-flowered Lipocarpha, 179 

Lithacodia albidula, 541 

Litholomia napaea, 542 

Lithomoia germana, 542 

Lithophane innominata, 542 

petulca, 542 

Lobophora nivigerata, 539 

Lobster, Atlantic, 82 

Locoweed, Field, 574 


_ Lolium perenne, 170,174 


Lomatia hirsuta, 132 
Longcore, J.R. and D.G. McAuley. Extraordinary Size and 
Survival of American Black Duck, Anas rubripes, 
Broods, 129 
Lonicera hispidula, 176 
Lontra canadensis, 96 
Loon, 210 
Common, 210 
Pacific, 210 
Red-throated, 256 
Loon (Gavia spp.) and Red-necked Grebe (Podiceps grise- 
gena) Populations in the Lower Matanuska-Susitna 
Valley, Alaska, Changes in, 210 
Lophocampa maculata, 540 
Lophodytes cucullatus, 131 
Lotus corniculatus, 365 


INDEX TO VOLUME 118 


663 


Lousewort, Oeder’s, 576 
Lupinus nootkatensis, 559 
Lutra canadensis, 267 
Luxilus cornutus, 523 
Luzula arcuata ssp. unalaschkensis, 559 
Lycaenidae, 537 
Lycia ursaria, 537 
Lycopodium complanatum, 196 
Lymantriidae, 540 
Lynx, 57,96,134 
Canada, 191,583 
Lynx canadensis, 57,96,191,583 
lynx, 134 
rufus, 40 
Lynx canadensis, Distributions, A Snow-tracking Protocol 
Used to Delineate Local Lynx, 583 
Lynx canadensis, in the Southern Canadian Rocky Mountains, 
Predation on Two Mule Deer, Odocoileus hemionus, 
by a Canada Lynx, 191 
Lynx, Lynx canadensis, Distributions, A Snow-tracking Pro- 
tocol Used to Delineate Local, 583 
Lynx, Lynx canadensis, in the Southern Canadian Rocky 
Mountains, Predation on Two Mule Deer, Odocoileus 
hemionus, by a Canada, 191 
Lyonetia prunifoliella, 533 
Lyonetiidae, 533 


Macaria bitactata, 537 
brunneata, 537 
loricaria, 537 
signaria, 537 
ulsterata, 537 
MacDonald, P., 444 
MacDonald, S.O., A.M. Runck, and J.A. Cook. The Heather 
Vole, Genus Phenacomys, in Alaska, 438 
MacDonnell, L., 444 
Macleaya cordata, 614 
Madden, E.M., 400 
Mahonia aquifolium, 176 
Maianthemum, 500 
canadense, 196,362 
canadense ssp. interius, 559 
Maier, T.J., 400 
Maitén, 132 
Malachigan, 523 
Malacosoma disstria, 540 
Mallard, 130 
Mallotus villosus, 262 
Malus sp., 125 
domestica, 299 
pumila, 361 
Manitoba*, Density and Survival of Lady Beetles (Coccinel- 
lidae) in Overwintering Sites in, 309 
Manitoba, Effects of Lodgepole Pine Dwarf Mistletoe, Arceu- 
thobium americanum, on Jack Pine, Pinus banksiana, 
Growth in, 595 
Manitoba, Hummock Vegetation at the Arctic Tree-line near 
Churchill, 590 
Manitoba, Ruby-throated Hummingbird, Archilochus colubris, 
Entanglements in Burdock, Arctium spp., at Delta 
Marsh, 85 
Manning (1911-1998), A Tribute to Thomas Henry, 618 
Maple, Manitoba, 309 
Mountain, 362 
Red, 41,96,251,277,361,508 


664 


Silver, 125 
Sugar, 41,124,251,361,507 
Marigane noire, 523 
Marine Turtle Newsletter, 156,294,484,645 
Marmot, Hoary, 440 
Vancouver, 403 
Marmota caligata, 438 
vancouverensis, 403 
Marten, 58,63 
American, 96,438 
Pine, 243 
Martes americana, 58,63,96,243,438 
pennanti, 56,96,583 
Martin, K., 579 
Maskinongé, 523 
Matteuccia struthiopteris, 46,253 
Maytenus boaria, 132 
McAlpine, D.F., S.A. Orchard, K.A. Sendall, and R. Palm. 
Status of marine turtles in British Columbia waters: 
a reassessment, 72 
McAuley, D.G., 129 
McCarthy, J, Reviews by, 150,152,640 
McIntyre, R.T., 115 
McKelvey, K.S., 583 
McKnight, J., L. Thomas, and D.L. Amirault. Piping Plover, 
Charadrius melodus, egg viability after seawater im- 
mersion, 448 
McLeish, W.J., 229 
McLoughlin, P.D., 239 
McNicholl, M.K., Review by, 285 
Meadow-foxtail, Carolina, 170 
Measures, L., B. Roberge, and R. Sears. Stranding of a Pygmy 
Sperm Whale, Kogia breviceps, in the Northern Gulf 
of St. Lawrence, Canada, 495 
Mech, L.D. and S. Tracy. Record High Wolf, Canis lupus, 
Pack Density, 127 
Mech, L.D., R.T. McIntyre and D.W. Smith. Unusual Beha- 
vior by Bison, Bison bison, Toward Elk, Cervus ela- 
phus, and Wolves, Canis lupus, 115 
Medicago sativa, 361 
Megasema c-nigrum, 543 
Melanchra adjuncta, 542 
Melanitta perspicillata, 264 
Melanitta perspicillata, in Coastal Southwestern British 
Columbia, Observations of Above-Surface Littoral 
Foraging in Two Sea Ducks, Barrow’s Goldeneye, 
Bucephala islandica, and Surf Scoter, 264 
Meleagris gallopavo silvestris, 251 
Meleagris gallopavo silvestris, Behavior in Central Ontario 
During Winter, Wild Turkey, 251 
Melica subulata, 176 
Méné a nageoires rouges, 523 
émeraude, 523 
jaune, 523 
Meningeal Worm, 40 
Mephitis mephitis, 40,69 
Merganser, Hooded, 131 
Mertensia paniculata var. alaskana, 559 
Mesoleuca ruficillata, 539 
Messier, F., 239 
Metalepsis salicarum, 543 
Metanema determinata, 538 
inatomaria, 538 
Meunier noir, 523 
Meyer, T.M., 185 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Mice, Peromyscus maniculatus, and Yellow-pine Chipmunks, 
Tamias amoenus, in Old Field and Orchard Habitats, | 
Population Dynamics of Deer, 299 

Micranthes kumlienti, 326 

nivalis, 326 
tenuis, 326 
Micropterus dolomieu, 523 
salmoides, 113,523 
Microtus, 345 
longicaudus, 300 
montanus, 300 
ochrogaster, 342 
pennsylvanicus, 342,400 

Milfoil, Eurasian, 318 

Millipede, 388 

Mills, T.K. and B.A. Andres. Changes in Loon (Gavia spp.) 
and Red-necked Grebe (Podiceps grisegena) Popula- 
tions in the Lower Matanuska-Susitna Valley, Alaska, 
210 

Millspaugh, J.J., B.E. Washburn, T.M. Meyer, J. Beringer, 
and L.P. Hansen. Immobilization of Clover-trapped 
White-tailed Deer, Odocoileus virginianus, with Med- 
etomidine and Ketamine, and Antagonism with Ati- 
pamezole, 185 

Mimulus guttatus, 559 

Mink, 96 

European, 185 
Minnow, Bluntnose, 318 
Minuartia dawsonensis, 559 

elegans, 559 

macrocarpa, 559 

obtusiloba, 166 

yukonensis, 559 

Minutes of the 125" Annual Business Meeting of the Ottawa 
Field-Naturalists’ Club 13 January 2004, 486 

Mistletoe, Lodgepole Pine Dwarf, 595 

Mistletoe, Arceuthobium americanum, on Jack Pine, Pinus 
banksiana, Growth in Manitoba, Effects of Lodgepole 
Pine Dwarf, 595 

Mizuhopecten yessoensis, 78 

Mniotilta varia, 230 

Molothrus ater, 382 

Mompha sp., 533 

albapalpella, 533 

terminella, 533 

Monarda punctata, 277 

Monkeyflower, Yellow, 576 

Monopis laevigella, 532 

spilotella, 532 

Montia fontana, 559 

Moore, J.W. and G.J. Kenagy. Consumption of Shrews, Sorex 
spp., by Arctic Grayling, Thymallus arcticus, 111 

Moose, 40,57,61,96,134,185,248,368,606 

Moose Sickness, 40 

Morrissey, C.A. and R.J. Olenick. American Dipper, Cinclus 
mexicanus, Preys Upon Larval Tailed Frogs, Ascaphus 
truei, 446 

Moss, 253 

Golden Fuzzy Fen, 591 

Irish, 226 

Stair-step, 591 

Moth, 388 

Moths and Butterflies (Lepidoptera) of the Boreal Mixedwood 
Forest near Lac La Biche, Alberta, Including New 
Provincial Records, 530 


2004 


| Mountain-heather, Yellow, 575 


Mountain-parsley, 560 


| Mouse, 342 


Deer, 299,342,402 
Great Basin Pocket, 300 
Meadow Jumping, 403 
Western Harvest, 300 
White-footed, 299,342 
Moxostoma spp., 521 
anisurum, 523 
carinatum, 523 
erythrurum, 526 
macrolepidotum, 523 
Mulders, R., 239 


| Mulinum spinosum, 132 
! Mullein, Great, 300 


Muntz, E.M. and B.R. Patterson. Evidence for the Use of 
Vocalization to Coordinate the Killing of a White- 
Tailed Deer, Odocoileus virginianus, by Coyotes, 
Canis latrans, 278 

Murray, D.F., Review by, 637 


' Murre, Common, 257 


Thick-billed, 602 
Musaraigne cendrée, 441 
fuligineuse, 441 
Musaraigne fuligineuse, Sorex fumeus, dans le nord-est du 
Québec, Extension de |’ aire de distribution connue de 
la, 441 


_ Muskrat, 267,602,607 
| Mussel, Bay, 264 
- Mustard, Tansy, 571 


Mustela spp., 96,306 
erminea, 243 
lutreola, 185 
nivalis, 403 
vison, 96 


| Mya arenaria, 78 


Myelopsis subtetricella, 536 


_ Myosotis scorpioides, 560 
| Myriophyllum spicatum, 318 


verticillatum, 559 
Myrrhis odorata, 614 
Myrrhis, Scented, 614 
Mythimna oxygala, 542 


_ Mytilicola orientalis, 77 
| Mytilus edulis, 264 


trossulus, 264 


Nadata gibbosa, 540 


| Naemia serriata, 227 


Naseux des rapides, 523 

Navarretia squarrosa, 169 

Neale, G.K., 95 

Nectria coecinea vat. faginata, 41 

Needle-and-thread, 564 

Needlegrass, Spreading, 564 

Nemapogon acapnopennella, 532 
roburella, 532 

Nematocampa resistaria, 538 

Nemophila parviflora, 176 

Nemophila, Small-flowered, 176 

Neneo, 132 

Neogobius melanostomus, 318 


INDEX TO VOLUME 118 


665 


Neogobius melanostomus Pallas (Gobiidae), and Ecotone 
Utilisation in St. Clair Lowland Waters, Ontario, Des- 
cription of Age-O Round Goby, 318 

Neotelphusa praefixa, 534 

Nepticulidae, 532 

Nerodia sipedon sipedon, 435 

Nerodia sipedon sipedon, Frequency of Tail Breakage of the 
Northern Watersnake, 435 

Nettle, Stinging, 567 

Neufeld, T., 605 

Newbury, T.L., 267 

Nguyen, L.P., J. Hamr, and G.H. Parker. Wild Turkey, Melea- 

gris gallopavo silvestris, Behavior in Central Ontario 

During Winter, 251 

Nicholson, P.H., 243 

Niditinea orleansella, 532 

Nites betulella, 533 

Noctuidae, 540 

Nostoc sp., 594 

Notemigonus crysoleucas, 523 

Nothofagus antarctica, 132 

pumilio, 132 

Notocelia culminana, 535 

Notodonta simplaria, 540 

Notodontidae, 540 

Notropis atherinoides, 319,523 

hudsonius, 318,523 

Noturus insignis, 523 

Nova Scotia, Continuing Environmental Change: An Example 

from, 39 

Nuphar polysepalum, 559 

Nuthatch, Red-breasted, 384 

Nycteola frigidana, 540 

Nymphalidae, 537 

Nymphalis antiopa, 537 

Nire, 132 


Oak, 342 
Bur, 310 
California Black, 383 
English, 125 
Gambel, 383 
Garry, 174,376 
Red, 124,251 
Oak Habitat in Southwestern British Columbia, Bird Com- 
munities of the Garry, 376 
Oat, 361,562 
Oatgrass, California, 169 
Oceanodroma leucorhoa, 258 
Oceanspray, 176 
Ochrolechia subplicans, 411 
subplicans ssp. hultenii, 420 
subplicans ssp. subplicans, 420 
Odocoileus spp., 368 
hemionus, 132,162,185,191,370,606 
hemionus columbianus, 162 
virginianus, 40,96, 127,185,278,360,370.606 
Odocoileus hemionus, by a Canada Lynx, Lynx canadensis, 
in the Southern Canadian Rocky Mountains, Preda- 
tion on Two Mule Deer, 191 
Odocoileus virginianus, by Coyotes, Canis latrans, Evidence 
for the Use of Vocalization to Coordinate the Killing 
of a White-Tailed Deer, 278 


666 


Odocoileus virginianus, Populations Living at Low and High 
Density in Southern Québec, Summer Diet of Two 
White-tailed Deer, 360 

Odocoileus virginianus, with Medetomidine and Ketamine, 
and Antagonism with Atipamezole, Immobilization 
of Clover-trapped White-tailed Deer, 185 

Oecophoridae, 533 

Oenothera biennis, 253 

Olenick, R.J., 446 

Olethreutes glaciana, 535 

metallicana, 535 
Olethreutinae, 535 
Oligia illocata, 541 

mactata, 541 

Oligochaete, 78 

Omisco, 523 

Oncorhynchus gorbuscha, 442 

keta, 248 

nerka, 111,249,443 

Oncorhynchus gorbuscha, Spawning Runs, Gulls, Larus spp.. 
Foraging at Pink Salmon, 442 

Ondatra zibethicus, 267,602,607 

Onion, Wild, 567 

Oniongrass, Alaska, 176 

Onoclea sensibilis, 253 

Ontario Bird Species, Predicting the Effects of Cerulean War- 
bler, Dendroica cerulea Management on Eastern, 229 

Ontario, Demographic patterns and limitations of Grey Wolves, 
Canis lupus, in and near Pukaskwa National Park, 95 

Ontario, Description of Age-O Round Goby, Neogobius mela- 
nostomus Pallas (Gobiidae), and Ecotone Utilisation 
in St. Clair Lowland Waters, 318 

Ontario During Winter, Wild Turkey, Meleagris gallopavo 
silvestris, Behavior in Central, 251 

Ontario, Effect of Fire Intensity and Depth of Burn on Low- 
bush Blueberry, Vaccinium angustifolium, and Velvet 
Leaf Blueberry, Vaccinium myrtilloides, Production 
in Eastern, 195 

Ontario Natural Heritage Information Centre Science and 
Information Newsletter 9(1) Winter 2004, 156 

Ontario: rabies management implications, Seasonal home 
ranges of raccoons using a common feeding site in rural 
eastern, 65 

Ontario, Significant Vascular Plant Records from the Hamil- 
ton Area, 612 

Ontario, The Northern True Katydid, Prerophylla camellifolia 
(Orthoptera: Pseudophyllidae), at Ottawa, 124 

Opegrapha sp., 411 

gyrocarpa, 406 
Operophtera bruceata, 539 
Oplopanax, 500 

horridum, 500 

Orache, Alaskan, 568 

Orchard, S.A., 72 

Oreamnos americanus, 370 

Oregon-grape, Tall, 176 

Oreodytes laevis, 427 

sanmarkii, 427 

Oreta rosea, 537 

Oriole, Baltimore, 232 

Orobanche fasciculata, 559 

Orthofidonia exornata, 537 

Orthosia hibisci, 542 

revicta, 542 

segregata, 542 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Oryx gazella, 185 
Osprey, 267 
Otisorex, 400 
Ottawa Field-Naturalists’ Club Awards for 2003, The, 646 
Otter, K.A., 354 
Otter, River, 96,267 
Sea, 270 
Otter, Enhydra lutris, Sightings off Haida Gwaii/Queen Char- - | 
lotte Islands, British Columbia, 1972-2002, Sea, 270 } 
Ouellet, J.-P., 360 
Ouitouche, 523 
Ovenbird, 232 
Ovis canadensis, 370 
Owl, Barred, 215,273 
Great Horned, 222,273 
Owl, Strix varia in Alberta: Distribution and Status, The — | 
Barred, 215 | 
Owl, Strix varia, in Labrador, First Record of a Barred, 273 
Oxytrope, Arctic, 574 
Oxytropis arctica, 558 
campestris ssp. varians, 559 
Oxyura jamaicensis, 604 
Oyster, Pacific, 77 


Pachnobia imperita, 543 
mixta, 543 
Palm, R., 72 
Palmaria palmata, 226 
Palthis angulalis, 540. 
Pandemis canadana, 535 
Pandion haliaetus, 267 
Panic-grass, Yellow, 197 
Panicum xanthophysum, 197 
Panthera tigris, 185 
Papaver croceum, 558 
nudicaule ssp. nudicaule, 558 
Papilio canadensis, 537 
Papilionidae, 537 
Paquet, P.C., 95 
Parapoynx maculalis, 537 
Parastichtis suspecta, 541 
Pardosa lapidicina, 123 
moesta, 122 
sternalis, 122 
Pardosa moesta and Pardosa sternalis (Araneae, Lycosidae) 1n 
Southeastern Idaho, Occurrence of Parasitoid Wasps, 
Baeus sp. and Gelis sp., in the Egg Sacs of the Wolf 
Spiders, 122 
Pardosa sternalis (Araneae, Lycosidae) in Southeastern Idaho, 
Occurrence of Parasitoid Wasps, Baeus sp. and Gelis 
sp., in the Egg Sacs of the Wolf Spiders Pardosa 
moesta and, 122 
Parectopa pennsylvaniella, 532 
Parelaphostrongylus tenuis, 40 
Pariseau, R., 521 
Parker, G.H., 251 
Parmelia saxatilis, 406 
Parornix conspicuella, 532 | 
Parrya arctica, 560 | 
Parsnip, Cow 196 
Partridge, S.T., 247 
Passerina cyanea, 232 
Patagonia, Observations of Interactions between Puma, Puma _ 
concolor, and Introduced European Red Deer, Cervus 
elaphus, in, 132 


— 


——S—= 


2004 


Patinopecten sp., 82 
| caurinus, 78 
Patterson, B.R., 278 
Pediasia dorsipunctella, 537 
| Pedicularis oederi, 559 
_Pelonaia corrugata, 79 
Penny, J.L., 174 


#))), Penry, L.B., 201 


| Pepper-grass, Common, 572 
| Perca flavescens, 521 

, Perchaude, 523 

| Percidae, 521 

| Percina caprodes, 523 

| copelandi, 523 


| Percopsidae, 523 

«  Percopsis omiscomaycus, 523 
| Perisoreus canadensis, 273 

| Perispasta caeculalis, 537 


_Perizoma basaliata, 539 


| Pero honestaria, 538 


| Perognathus parvus, 300 

Peromyscus spp., 342 

leucopus, 299,342 

| maniculatus, 299,342,402 

| Peromyscus maniculatus, and Yellow-pine Chipmunks, Tamias 
| amoenus, in Old Field and Orchard Habitats, Popu- 
| lation Dynamics of Deer Mice, 299 


| Perry, N.D., D.T. Stewart, EM. Madden, and T.J. Maier. New 
| Records for the Arctic Shrew, Sorex arcticus and the 
i] Newly Recognized Maritime Shrew, Sorex mariti- 


mensis, 400 
| Peterkin, P., Review by, 641 
| Petrel, Leach’s Storm, 258 


| Petromyzontidae, 523 
| Phacelia, Macbryde’s, 576 
| Phacelia mollis, 559 


Phalacrocorax auritus, 258 
carbo, 258 


j Phalaenophana pyramusalis, 540 


Phalaenostola hanhami, 540 


| Phaneta awemeana, 535 


convergana, 535 

parmatana, 535 
Phenacomys intermedius, 438 

intermedius intermedius, 438 

ungava, 438 


i Phenacomys, in Alaska, The Heather Vole, Genus, 438 


Pheosia rimosa, 540 

Pheucticus ludovicianus, 232 

Phillips, F., 273 

Phleum pratense, 253 

Phlogophora periculosa, 542 

Phlox hoodii, 559 

Phlox, Moss, 575 

Phlyctaenia coronata, 537 

Phoca hispida, 395,602 

Phragmatobia assimilans, 540 

Phyllodesma americana, 539 

Phyllodoce glanduliflora, 559 

Phyllonorycter sp., 532 
martiella, 532 

Physcia caesia, 411 

Physeter macrocephalus, 495 

Picard, [., 441 

Picea spp., 57,61 


INDEX TO VOLUME 118 


667 


engelmannii, 191,584 
glauca, 46,95,211,218,249,251,274,361,370,388,401, 
504,530,590 
glauca X engelmannii, 355 
mariana, 62,95,268,274,370,388,504,590 
rubens, 41,52,510 
sitchensis, 499 
Picoides pubescens, 358 
Pieridae, 537 
Pieris oleracea, 537 
Pimephales notatus, 318 
Pine, Eastern White, 251 
Ground, 196 
Jack, 95,196,253,388,595 
Lodgepole, 58,62,191,355,370,499,561,584,595 
Ponderosa, 198,300,383,595 
Red, 196,251 
White, 41,196,277 
Whitebark, 584 
Pine, Pinus banksiana, Growth in Manitoba, Effects of Lodge- 
pole Pine Dwarf Mistletoe, Arceuthobium america- 
num, on Jack, 595 
Pinguicula villosa, 559 
Pinus spp., 60 
albicaulis, 584 
banksiana, 95,196,253,388,595 
contorta, 58,62,191,370,499,584,595 
contorta ssp. latifolia, 559 
contorta var. latifolia, 355 
ponderosa, 198,300,383,595 
resinosa, 196,251 
strobus, 41,196,251,277 
Pinus banksiana, Growth in Manitoba, Effects of Lodgepole 
Pine Dwarf Mistletoe, Arceuthobium americanum, on 
Jack Pine, 595 
Pipilo maculatus, 378 
Piranga ludoviciana, 378 
Piranga olivacea, 232 
Pisaster ochraceus, 265 
Pissodes strobi, 41 
Pitcher, K., 270 
Placopsis gelida, 424 
lambii, 406 
Plagiobothrys scouleri, 170 
Plagodis alcoolaria, 538 
phlogosaria, 538 
pulveraria, 538 
Plaintain, Seaside, 577 
Plantago spp., 247 
canescens, 559 
maritima, 559 
Plasmodium spp., 237 
Platarctia parthenos, 540 
Platichthys stellatus, 248 
Platynota idaeusalis, 535 
Platyphyllum concavum, 125 
Platypolia anceps, 542 
Plemyria georgii, 538 
Pleurozium schreberi, 591 
Plover, Piping, 444.448 
Snowy, 445 
Plover, Charadrius melodus, egg viability after seawater im- 
mersion, Piping, 448 
Plover, Charadrius melodus, Nesting 
Sequential Polyandry in Piping, 


in Eastern Canada, 
444 


668 


Plume-poppy, 614 
Plusia putnami, 541 
Plusiinae, 541 
Plutella xylostella, 533 
Plutellidae, 533 
Poa spp., 300 
annua, 560 
compressa, 277 
interior, 558 
leptocoma, 559 
nemoralis, 558 
pratensis, 176 
secunda, 559 
Podiceps grisegena, 210 
Podiceps grisegena) Populations in the Lower Matanuska- 
Susitna Valley, Alaska, Changes in Loon (Gavia spp.) 
and Red-necked Grebe, 210 
Podistera macounii, 559 
Poecile atricapillus, 232 
hudsonicus, 48 
rufescens, 378 
Pohl, G.R., D.W. Langor, J.-F. Landry, and J.R. Spence. Moths 
and Butterflies (Lepidoptera) of the Boreal Mixed- 
wood Forest near Lac La Biche, Alberta, Including 
New Provincial Records, 530 
Point Pelee Natural History News 3(4), 155 
Polemonium acutiflorum forma lacteum, 558 
boreale forma albiflorum, 558 
Polia imbrifera, 542 
nimbosa, 542 
Polioptila caerulea, 232 
Polix coloradella, 533 
Polychidium muscicola, 424 
Polychrysia esmerelda, 541 
Polygonatum pubescens, 196 
Polygonia satyrus, 537 
Polygonum lapathifolium, 559 
viviparum, 591 
Polystichum kruckebergii, 164 
lemmonii, 164 
mohrioides, 164 
scopulinum, 164 
Polystichum lemmonii, a Threatened Fern in Canada, Con- 
servation Evaluation of Lemmon’s Holly Fern, 164 
Polytrichadelphus lyellii, 421 
Pomoxis nigromaculatus, 523 
Pond-lily, Yellow, 569 
Pondweed, 318 
Curly, 318 
Flatstemmed, 561 
Sago, 561 
Variable-leaved, 561 
White-stemmed, 561 
Popcornflower, Scouler’s, 170 
Poplar, Balsam, 62,218,268,530 
Populus spp., 52,57,61,511 
balsamifera, 62,136,218,268,530 
balsamifera ssp. trichocarpa, 355 
deltoides, 309 
grandidentata, 136 
tremuloides, 46,62,96,136,211,218,251,268,274,277, 
316,342,355,361,370,388,5 15,530,584 
trichocarpa, 211 
Porbeagle, 245 
Porcupine, 607 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Porina chlorotica, 421 
pacifica, 406 
Porpidea speirea, 415 
carlottiana, 411 
contraponenda, 406 
speirea, 406 
thomsonii, 406 
Poszig, D., C.D. Apps, and A. Dibb. Predation on Two Mule » 
Deer, Odocoileus hemionus, by a Canada Lynx, Lynx : 
canadensis, in the Southern Canadian Rocky Moun- 
tains, 191 
Potamogeton spp., 318,365 
alpinus ssp. tenuifolius, 559 
berchtoldii, 559 
crispus, 318 
gramineus, 559 
pectinatus, 559 
praelongus, 559 
subsibiricus, 559 
zosteriformis, 559 
Potentilla arguta ssp. convallaria, 573 
arguta var. convallaria, 559 
Prairie-crocus, 569 
Prest (1913-2003), A Tribute to Victor Kent, 626 
Primrose, Evening, 253 
Primula eximia, 559 
tschuktschorum ssp. cairnesiana, 575 
Prince Edward Island, A Mapping of the Present and Past 
Forest-types of, 504 
Prince Edward Island, Use of Eelgrass, Zostera marina, Wrack 
by Three Species of Ladybird Beetles (Coleoptera: 
Coccinellidae) in, 225 
Prionoxystus robiniae, 534 
Procyon lotor, 65 
Prodoxidae, 532 
Propylea quatuordecimpunctata, 225 
Proterorhinus marmoratus, 318 
Protitame virginalis, 537 
Protoboarmia porcelaria, 537 
Protolampra rufipectus, 543 
Protolithocolletis lathyri, 532 
Protorthodes oviduca, 542 
Protothaca staminea, 77 
Pruitt, W.O., Jr., Review by, 638 
Prunus sp., 136 
pennsylvanica, 365 
pumila, 196 
serotina, 365 
virginiana, 253,362 
Pseudacris maculata, 136 
Pseudexentera oregona, 536 
Pseudohermonassa tenuicula, 543 
Pseudosciaphila duplex, 535 
Pseudothyatira cymatophoroides, 537 
Pseudotsuga menziesii, 159,376,584 
menziesii var. glauca, 355 
menziesii var. menziesii, 599 
Psilocarphus elatior, 169 
tenellus var. tenellus, 169 
Psilocarphus elatior, an Endangered Herb in Canada, Con- 
servation Evaluation of the Pacific Population of Tall 
Woolly-heads, 169 
Psyllobora virginimaculata, 312 
Ptarmigan, Willow, 602 
Pteridium aquilinum, 196,251 


ieee 


2004 


' Pterophoridae, 536 
Pterophylla, 125 
camellifolia, 124 
Pterophylla camellifolia (Orthoptera: Pseudophyllidae), at 
Ottawa, Ontario, The Northern True Katydid, 124 
Ptychoramphus aleuticus, 616 
| Puccinellia interior, 559 


| Puffin, Atlantic, 257,604,615 


Horned, 615 
| Tufted, 615 
| Pulsatilla ludoviciana, 559 
Puma concolor, 132,605 
concolor vancouverensis, 159 
|Puma, 132 
| Puma concolor, and Introduced European Red Deer, Cervus 
elaphus, in Patagonia, Observations of Interactions 
between Puma, 132 
‘Puma concolor, in Elk Island National Park, Alberta, First 
Record of Mountain Lions, 605 


Puma concolor vancouverensis, Morphology and Population 


| 


Characteristics of Vancouver Island Cougars, 159 
Puma, Puma concolor, and Introduced European Red Deer, 
Cervus elaphus, in Patagonia, Observations of Inter- 
| actions between, 132 
| Punctelia stictica, 424 
| Pyralidae, 536 
! Pyrausta borealis, 537 
nicalis, 537 
| Pyrenocollema halodytes, 410 


| sublitorale, 424 


Pyrola minor, 559 


Quackgrass, 253 


: Quail, California, 378 


Québec, Extension de I’aire de distribution connue de la Mus- 
araigne fuligineuse, Sorex fumeus, dans le nord-est du, 
44] 


Quebec, Forked Three-awned Grass, Aristida basiramea 


Engelm. ex Vasey: A New Addition to the Flora of, 

276 

Québec, Inventaire printanier d’une frayére multispécifique: 

Vichtyofaune des rapides de la riviére Gatineau, 521 

Québec, Premiéres mentions de la Couleuvre mince, Tham- 

nophis sauritus septentrionalis, au, 135 

Québec, Summer Diet of Two White-tailed Deer, Odocoileus 
virginianus, Populations Living at Low and High Den- 


sity in Southern, 360 
—Quenouilles, 136 


Quercus spp., 342,365 
alba, 342 
candicans, 383 
gambellii, 383 
garryana, 174,376 
kelloggii, 383 
laurina, 383 
macrocarpa, 310,342 


i obtusata, 383 


robur, 125 
rubra, 124,251,342 
rugosa, 383 


' Queue 4 tache noire, 523 
- Quillwort, Bristle-like, 560 


Maritime, 560 


INDEX TO VOLUME 


118 669 


Raccoon, 65 
Raccoons using a common feeding site in rural eastern On- 
tario: rabies management implications, Seasonal home 
ranges of, 65 
Radal, 132 
Ragweed, Hybrid, 613 
Ragwort, Dryland, 577 
Rail, J.-F. and G. Chapdelaine. Fifteenth Census of Seabird 
Populations in the Sanctuaries of the North Shore of 
the Gulf of St. Lawrence, 1998-1999, 256 
Rainette faux-grillon boréale, 136 
Raisin, Northern Wild, 253 
Ramalina siliquosa, 420 
Rana aurora, 448 
Randa, L.A. and J.A. Yunger. The Influence of Prey Avail- 
ability and Vegetation Characteristics on Scent Sta- 
tion Visitation Rates of Coyotes, Canis latrans, in a 
Heterogeneous Environment, 341 
Rangifer tarandus, 39,119,192,241,362,602 
tarandus caribou, 57,61,119,368 
tarandus tarandus, 96,185 
Rangifer tarandus caribou, in Saskatchewan, Morphology 
of Female Woodland Caribou, 119 
Ranunculus aquatilis var. subrigidus, 559 
occidentalis var. brevistylis, 559 
Raphia frater, 541 
Raptor, 306 
Raseux-de-terre gris, 523 
noir, 523 
Raspberry, 251 
Raum-Suryan, K., K. Pitcher, and R. Lamy. Sea Otter, Enhy- 
dra lutris, Sightings off Haida Gwaii/Queen Charlotte 
Islands, British Columbia, 1972-2002, 270 
Raven, 273 
Razorbill, 257 
Redcedar, Western, 159 
Redstart, American, 232 
Regulus calendula, 48,86 
satrapa, 48,86 
Reindeer, 185 
Reithrodontomys megalotis, 300 
Renaud, C.B., 521 
RENEW. 2004. Recovery of Nationally Endangered Wildlife 
in Canada. Annual Report (14), 644 
Retinia burkeana, 535 
Rettie, W.J. Morphoiogy of Female Woodland Caribou, 
Rangifer tarandus caribou, in Saskatchewan, 119 
Rheum rhabarbarum, 560 
Rhigognostis interrupta, 533 
Rhinanathus minor ssp. borealis, 559 
cataractae, 523 
Rhizocarpon, 411 
cinereovirens, 421 
geminatum, 411 
haidensis, 411 
hensseniae, 408 
hochstetteri, 411 
maritimum, 421 
Rhubarb, 568 
Ribes, 500 
oxyacanthoides ssp. oxyacanthoides, 559 
Richardson, E.S. and R.K. Brook. Excavation of an Arctic Fox, 
Alopex lagopus, den by a Polar Bear, Ursus maritimus, 
602 
Rinodina gennari, 408 


670 


Rissa tridactyla, 258 
Rivula propinqualis, 540 
Rivulinae, 540 
Roberge, B., 495 
Robertson, R.J., 229 
Robin, American, 381 
Rock-Cress, Wall, 570 
Rockweed, 406 
Rorippa barbareifolia, 558 
Rosa blanda, 253 
woodsii, 559 
Rosatte, R.C., 65 
Rose, Smooth Wild, 253 
Western, 574 
Rosell, F., 434 
Rothfels, C. Significant Vascular Plant Records from the Ham- 
ilton Area, Ontario, 612 
Rubus spp., 251,365,500 
ideaus, 197 
spectabilis, 500 
Ruggiero, L.F., 583 
Rumex crispus, 560 
Runck, A.M., 438 
Rush, 400 
Toad, 170,567 
Russell, 1904-1998, A Tribute to Loris Shano, 451 
Ryegrass, Perennial, 170,174 


Sabia conica, 78 
Sagebrush, 300 
Sagittaria sp., 318 
Salal, 159 
Salamander, Cope’s, 448 
Pacific Giant, 448 
Salamandre sombre des montagnes, 136 
Salix spp., 57,136,180,211,247,253,268,356,365,370,530 
alaxensis ssp. longistylis, 559 
amagdaloides, 309 
glauca var. acutifolia, 559 
novae-angliae, 567 
pseudomyrsinites, 559 
rotundifolia ssp. dodgeana, 559 
Salmo clarki, 448 
salar, 77 
trutta, 113,523 
Salmon, Atlantic, 77 
Chum, 248 
Pink, 442 
Red, 249 
Sockeye, 111,443 
Salmon, Oncorhynchus gorbuscha, Spawning Runs, Gulls, 
Larus spp., Foraging at Pink, 442 
Salmonidae, 523 
Salsify, Yellow, 300 
Salvelinus namaycush, 523 
Sambucus, 501 
racemosa, 500 
racemosa ssp. pubens, 559 
Samuel, W.M., 95 
Sander spp., 521 
canadensis, 523 
vitreus, 523 
Sander-Regier, R., Reviews by, 153,474,477,478,632 
Sandlance, 262 
Sandpiper, Spotted, 444 


THE CANADIAN FIELD-NATURALIST 


Vol. 118: 


Sandwort, Alpine, 166 
Sanicle, Pacific, 174 
Sanicula crassicaulis var. crassicaulis, 174 
Sargassum muticum, 77 
Sarrothripinae, 540 
Sarsasparilla, Wild, 46 
Saskatchewan, Morphology of Female Woodland Caribou, , 
Rangifer tarandus caribou, in, 119 
Saskatoon, 573 
Saxidomus giganteus, 77 
Saxifraga, 326 
caespitosa, 559 
eschscholtzii, 559 
foliolosa, 327,559 
gaspensis, 327 
hieracifolia, 336 
nelsoniana ssp. pacifica, 559 
nivalis, 326 
nivalis var. gaspensis, 327 
nivalis var. labradorica, 327 
nivalis var. rufopilosa, 327 
nivalis var. tenuis, 326 
rufopilosa, 327,559 
tenuis, 326 
tricuspidata, 559 
Saxifrage, Prickly, 573 
Scale, Woolly Beech, 41 
Scallop, Weathervane, 82 
Scaup, Greater, 268 
Schinus patagonicus, 132 
Schizoporella unicornis, 79 
Schizura leptinoides, 540 
unicornis, 540 
Schmelzer, [. and F. Phillips. First Record of a Barred Owl, 
Strix varia, in Labrador, 273 
Schmiegelow, F.K.A., 368 
Schoen, J.W., 247 
Sciaenidae, 523 
Scirpus spp., 318 
americanus, 226 
cyperinus, 402 
maritimus, 226 
rollandii, 559 
Scoparia biplagialis, 531 
Scopula ancellata, 538 
frigidaria, 538 
junctaria, 538 
limboundata, 538 
Scoter, Surf, 264 
Scoter, Melanitta perspicillata, in Coastal Southwestern 
British Columbia, Observations of Above-Surface 
Littoral Foraging in Two Sea Ducks, Barrow’s Gold- 
eneye, Bucephala islandica, and Surf, 264 
Scouring-rush, Variegated, 591 
Scymnus (Pullus) brullei, 312 
Seal, Ringed, 395,602 
Sealy, S.G., 85 
Sears, R., 495 
Sea Star, Ochre, 265 
Seaweed, 77 
Sargassum, 78 
Seburn, C., Reviews by, 149,635 
Seburn, D., Reviews by, 140,143,468,471 
Sedge, 318,400 
Blunt, 565 


=== 


Green-sheathed, 170 

Houton’s, 197 

Mud, 591 

One-sided, 169 

Ross’, 566 

Slender-beaked, 169 

Two-toned, 564 

i Water, 591 

Seiurus aurocapilla, 232 
noveboracensis, 232 

| Selasphorus rufus, 378 

| Selenia alciphearia, 538 

| Semibalanus spp., 406 

| Semioscopis inornata, 533 

Semotilus corporalis, 523 

| Sendall, K.A., 72 

Senecio eremophilus, 558 


| Sergeant, D.E., Review by, 144 

| Serviceberry, 196 

| Setaria viridis, 558 

| Setophaga ruticilla, 232 

| Seymour, K.L. A Tribute to Loris Shano Russell, 1904-1998, 


| 451 

| Shackleton, D.M., 159 
| Shad, American, 77 

| Gizzard, 318 


| Sharpe. F., 434 
| Shaverdo, H.V. and D.J. Giberson. Predaceous Water Beetles 


lected Along the Horton and Thelon Rivers in the 


(Coleoptera: Adephaga: Dytiscidae, Gyrinidae) Col- 
| Arctic Central Barrens of Canada, 425 


| Sheep, Bighorn, 370 


Shepherd, German, 247 


| Shepherd’s-purse, 571 


| Shiner, Emerald, 319 
| Spottail, 318 
| Shooting-star, Broad-leaved, 175 
| Shrew, 111 
| Arctic, 400 
| Gaspé, 403 
| Maritime, 400 
] 
\ 


Masked, 400 

Northern Short-tailed, 402 

Pygmy, 402 

Smoky, 400 

Tundra, 400 

| Shrew, Sorex arcticus and the Newly Recognized Maritime 


Shrew, Sorex maritimensis, New Records for the Arctic, 
400 


|| Shrew, Sorex maritimensis, New Records for the Arctic Shrew, 
Til 


Sorex arcticus and the Newly Recognized Maritime, 
400 

Shrews, Sorex spp., by Arctic Grayling, Thymallus arcticus, 
Consumption of, 111 

Sicya macularia, 538 

Silene involucrata ssp. tenella, 559 

vulgaris, 560 
| williamsii, 559 
Silverside, Brook, 319 

| Sinapis arvensis, 560 

| Sinoe sp., 534 

Siskin, Pine, 378 

| Sisymbrium altissimum, 300 


| Sitta canadensis, 384 


Skunk, Striped, 40,69 


INDEX TO VOLUME 118 


671 


Skunkweed, 169 
Sloan, N.A., 405 
Sloan, N.A. and P.M. Bartier. Introduced Marine Species in 
the Haida Gwaii (Queen Charlotte Islands) Region, 
British Columbia, 77 
Slug, 388 
Smartwood, Pale, 568 
Smerinthus cerisyi, 540 
Smith, C.M., Reviews by, 284,287 
Smith, D.W., 115 
Smith, T., Reviews by, 146,148,480,641 
Smith, T.S., S.T. Partridge, and J.W. Schoen. Interactions of 
Brown Bears, Ursus arctos, and Gray Wolves, Canis 
lupus, at Katmai National Park and Preserve, Alas- 
ka, 247 
Smith, T.W., G.W. Douglas and A.G. Harris. Conservation 
Evaluation of Small-flowered Lipocarpha, Lipocarpha 
micrantha (Cyperaceae), in Canada, 179 
Snail, 78,388 
Snake, Butler’s Garter, 435 
Eastern Garter, 435 
Garter, 448 
Northern Ribbon, 435 
Snowberry, Common, 175 
Sobey, D.G. and W.M. Glen. A Mapping of the Present and 
Past Forest-types of Prince Edward Island, 504 
Solidago spp., 253,365 
altissima, 342 
canadensis var. salebrosa, 560 
Solomon’s Seal, Hairy, 196 
Somateria mollissima, 258 
Sonchus sp., 365 
arvensis ssp. uliginosus, 560 
Sorbus americana, 362 
Sorex spp., 111,400 
araneus, 113 
arcticus, 400 
arcticus arcticus, 402 
arcticus laricorum, 402 
arcticus maritimensis, 400 
cinereus, 111,400,441 
fumeus, 400,441 
gaspensis, 403 
hoyi, 112,402 
maritimensis, 400 
monticolus, 111,438 
tundrensis, 112,400 
yukonicus, 112 
Sorex arcticus and the Newly Recognized Maritime Shrew, 
Sorex maritimensis, New Records for the Arctic Shrew, 
400 
Sorex fumeus, dans le nord-est du Québec, Extension de l’aire 
de distribution connue de la Musaraigne fuligineuse. 
44] 
Sorex maritimensis, New Records for the Arctic Shrew, Sorex 
arcticus and the Newly Recognized Maritime Shrew, 
400 
Sorex spp., by Arctic Grayling, Thymallus arcticus, Con- 
sumption of Shrews, 111 
Sorgastrum nutans, 342 
South Dakota, Spring Dispersal Patterns of Red-winged 
Blackbirds, Agelaius phoeniceus, Staging in East- 
ern, 201 
Sow-thistle, Perennial, 578 
Soybean, 253 


672 


Spargania luctuata, 539 
Sparganothis sp., 535 
reticulatana, 535 
xanthoides, 535 
Sparrow, Chipping, 378 
Song, 378 
White-crowned, 378 
Spartina spp., 400 
alterniflora, 226 
patens, 226 
Spence, J.R., 530 
Spermophilus spp., 90 
beecheyi, 90 
beldingi, 90 
brunneus, 91 
columbianus, 90 
parryii, 91 
richardsonii, 90 
tridecemlineatus, 90 
Spermophilus richardsonii, Multiple Mating Results in Mul- 
tiple Paternity in Richardson’s Ground Squirrels, 90 
Sphagnum, 499 
Sphingidae, 540 
Spider, Wolf, 122 
Spiders Pardosa moesta and Pardosa sternalis (Araneae, 
Lycosidae) in Southeastern Idaho, Occurrence of Par- 
asitoid Wasps, Baeus sp. and Gelis sp., in the Egg Sacs 
of the Wolf, 122 
Spilonema revertens, 410 
Spiraea latifolia, 253 
Spiranthes romanzoffiana, 560 
Spizella passerine, 378 
Springbeauty, Alpine, 568 
Springtail, 389 
Spruce, Black, 57,61,95,268,274,370,388,504,590 
Engelmann, 191,584 
Norway, 124 
Red, 41,52,514 
Sitka, 499 
White, 46,57,61,95,211,218,249,251,274,355,361,370, 
388,401,504,530,590 
Squid, 497 
Squires, J.R., K.S. McKelvey, and L.F. Ruggiero. A Snow- 
tracking Protocol Used to Delineate Local Lynx, Lynx 
canadensis, Distributions, 583 
Squirrel, Arctic Ground, 91 
Belding’s Ground, 90 
California Ground, 90 
Columbian Ground, 90 
Idaho Ground, 91 
Red, 607 
Richardson’s Ground, 90 
Thirteen-lined Ground, 90 
Squirrels, Spermophilus richardsonii, Multiple Mating Results 
in Multiple Paternity in Richardson’s Ground, 90 
Starflower, 575 
Starling, European, 378 
Stellaria umbellata, 560 
Stenolechia sp., 534 
Sterna caspia, 258 
hirundo, 258 
paradisaea, 258 
Sternotherus odoratus, 136 
Sterrhinae, 538 
Stethorus punctum, 312 


THE CANADIAN FIELD-NATURALIST 


Stewart, D.T., 400 
Stictotarsus griseostriatus, 430 
Stigmella sp., 532 
Stipa comata, 560 
nelsonii ssp. dorei, 560 
richardsonii, 560 
speciosa var. major, 132 
Stoffel, M.J., Review by, 284 
Storeria dekayi, 136 
Storm-Petrel, Leach’s, 259 
Strawberry-blite, 568 
Strepsimaninae, 540 
Streptopus, 500 
amplexifolius, 500 
Strix varia, 215,273 
Strix varia in Alberta: Distribution and Status, The Barred 
Owl, 215 | 
Strix varia, in Labrador, First Record of a Barred Owl, 273 
Sturnus vulgaris, 378 
Subularia aquatica ssp. americana, 560 
Sucker, White, 319 
Sullivan, D.S., 299 i 
Sullivan, T.P., D.S. Sullivan, and E.J. Hogue. Population Dy- 
namics of Deer Mice, Peromyscus maniculatus, an 
Yellow-pine Chipmunks, Jamias amoenus, in Old dl 
Field and Orchard Habitats, 299 | 
Swallow, Barn 230 i! 
Violet-green, 379 
Swammerdamia caesiella, 532 
Sweet, Meadow, 253 
Sylvilagus, 345 
floridanus, 342 
Symphoricarpos albus, 175 
Symphyotrichum laurentianum, 105 
Symphyotrichum laurentianum, (Fernald) Nesom, Estimation } 
of Seed Bank and Seed Viability of the Gulf of Saint | 
Lawrence Aster, 105 
Synaptomys borealis, 438 
Synclita obliteralis, 537 
Syncopacma sp., 534 
Syndemis afflictana, 535 
Syngrapha alias, 541 
octoscripta, 541 
rectangula, 541 
viridisigma, 541 


Takats Priestley, L. The Barred Owl, Strix varia in Alberta: 
Distribution and Status, 215 

Tamarack, 63,370,515,590 

Tamias amoenus, 299 

Tamias amoenus, in Old Field and Orchard Habitats, Popu- 
lation Dynamics of Deer Mice, Peromyscus manicu- 
latus, and Yellow-pine Chipmunks, 299 

Tamiasciurus hudsonicus, 607 

Tanacetum vulgare, 560 

Tanager, Scarlet, 232 

Western, 378 

Taniva albolineana, 535 

Tansey, Common, 578 

Taraxacum officinale, 365 

Tardif, J.C., 595 

Taxus canadensis, 41,365 

Tea, Common Labrador, 591 

Teal, Green-winged, 131 

Teleiodes proximella, 534 


IF 


1 


ek ITE 


2004 


SS 


Telfer, E.S. Continuing Environmental Change: An Example 
from Nova Scotia, 39 
| Tephromela atra, 424 
Tern, Arctic, 257 
Caspian, 258 
Common, 257 
Tews, J. Hummock Vegetation at the Arctic Tree-line near 
i} Churchill, Manitoba, 590 
Thamnophis spp., 448 
| butleri, 435 
sauritus, 135,435 
| sauritus septentrionalis, 135 
sirtalis, 435 
sirtalis sirtalis, 135 
Thamnophis sauritus septentrionalis, aa Québec, Premieres 
mentions de la Couleuvre mince, 135 

| Thistle, Swamp, 253 
Thlaspi arvense, 560 
Thomas, L., 448 

Thomsen, L.R., F. Sharpe, and F. Rosell. Collapsing Burrow 
| Causes Death of a Eurasian Beaver, Castor fiber, 434 
Thrush, Swainson’s, 382 
! Wood, 232 
| Thryomanes bewickii, 378 


‘f 


| Thuja occidentalis, 96,136,253,361,508 


| plicata, 159 
_Thymallus arcticus, 111 
thymallus, 113 
Thymallus arcticus, Consumption of Shrews, Sorex spp., by 
| Arctic Grayling, 111 
Tick, American Dog, 42 
Tiger, 185 
Tilia americana, 253,310 


_| Timmins, C.A., 318 


| Timothy, 253 

| Tineidae, 532 

| Tinline, R.R., 65 

| Todd, G., 90 

Tofieldia coccinea, 560 

| Tomenthypnum nitens, 591 

Tortricidae, 534 

| Tortricinae, 534 

| Tortue musquée, 136 

Totton, S.C., R.C. Rosatte, R.R. Tinline, and L.L. Bigler. Sea- 
sonal home ranges of raccoons using a common feed- 
ing site in rural eastern Ontario: rabies management 
implications, 65 

| Touladi, 523 

_Towhee, Spotted, 378 

| Tracheoniscus rathkei, 388 


i Tracy, S., 127 


Tragopogon dubius, 300 

| Trechus obtusus, 79 

| Trichodezia albovittata, 539 

| Trientalis europaea, 560 

Trifolium spp., 253,362 
dubium, 170 

| pratense, 560 

Triglochin spp., 247 
maritima, 226 
palustre, 560 

Trillium sp., 365 

Triphosa haesitata, 539 

|. Triteleia grandiflora, 174 

grandiflora var. howellii, 174 


INDEX TO VOLUME 118 


673 


howellti, 174 
Triteleia, Howell’s, 174 
Triteleia howellii, an Endangered Lily in Canada, Conserva- 
tion Evaluation of Howell’s Triteleia, 174 
Triteleia, Triteleia howellii, an Endangered Lily in Canada, 
Conservation Evaluation of Howell’s, 174 
Troglodytes aedon, 378 
Trout, Brown, 113 
Cutthroat, 448 
Truite brune, 523 
Trypanosoma, 236 
avium, 237 
Tsuga spp., 499 
canadensis, 41,251,361,508 
heterophylla, 159 
mertensiana, 159 
Tubularia crocea, 78 
Tumble-mustard, Tall, 300 
Tunicate, 79 
Turdus migratorius, 381 
Turkey, Eastern Wild, 251 
Turkey, Meleagris gallopavo silvestris, Behavior in Central 
Ontario During Winter, Wild, 251 
Turnock, W.J. and I.L. Wise. Density and Survival of Lady 
Beetles (Coccinellidae) in Overwintering Sites in 
Manitoba*, 309 
Turtle, Green, 72,579,611 
Hawksbill, 580 
Kemp’s Ridley, 580 
Leatherback, 72,579 
Loggerhead, 579 
Murray Short-necked, 610 
Snapping, 610 
Turtle, Caretta caretta, and the First Record of a Green Tur- 
tle in Atlantic Canada, Hybridization between a Green 
Turtle, Chelonia mydas, and Loggerhead, 579 
Turtle, Chelonia mydas, and Loggerhead Turtle, Caretta caret- 
ta, and the First Record of a Green Turtle in Atlantic 
Canada, Hybridization between a Green, 579 
Turtle in Atlantic Canada, Hybridization between a Green Tur- 
tle, Chelonia mydas, and Loggerhead Turtle, Caretta 
caretta, and the First Record of a Green, 579 
Turtles, Chelydra serpentina?, Is Cost of Locomotion the 
Reason for Prolonged Nesting Forays of Snapping. 
610 
Turtles in British Columbia waters: a reassessment, Status 
of marine, 72 
Tylothallia biformigera, 424 
Typha spp., 136,318 
latifolia, 253,546,560 


Udea itysalis, 537 
Ulmus americana, 310,508 
Underwood, T.J., 85 
Untereiner, W.A., 90 
Update on Quebec Amphibian and Reptile Atlas Project: Rana- 
Saura 6(2), 294 
Uraniidae, 539 
Uria aalge, 258 
lomvia, 602 
Urodidae, 536 
Ursus spp., 443 
americanus, 41.96,370 
arctos, 117,134,239,247,370,499,603 
maritimus, 395,602 


674 


Ursus arctos, and Gray Wolves, Canis lupus, at Katmai Nation- 
al Park and Preserve, Alaska, Interactions of Brown 
Bears, 247 

Ursus arctos, in Southeastern Alaska, Seed Dispersal by 
Brown Bears, 499 

Ursus arctos, in the Central Canadian Arctic, Movements of 
Subadult Male Grizzly Bears, 239 

Ursus maritimus, Excavation of an Arctic Fox, Alopex lago- 
pus, den by a Polar Bear, 602 

Ursus maritimus, in the Alaskan Beaufort, Chukchi, and 
Northern Bering Seas, Observations of Habitat Use 
by Polar Bears, 395 

Urtica dioica ssp. gracilis, 560 

Utricularia minor, 560 


Vaccinium sp., 159,251,500 
alaskense, 500 
angustifolium, 195 
globulare, 198 
membranaceum, 560 
myrtilloides, 195 
ovalifolium, 500 
vitis-idaea, 591 
Vaccinium angustifolium, and Velvet Leaf Blueberry, Vac- 
cinium myrtilloides, Production in Eastern Ontario, 
Effect of Fire Intensity and Depth of Burn on Low- 
bush Blueberry, 195 
Vaccinium myrtilloides, Production in Eastern Ontario, Effect 
of Fire Intensity and Depth of Burn on Lowbush Blue- 
berry, Vaccinium angustifolium, and Velvet Leaf Blue- 
berry, 195 
Vahlodea atropurpurea, 560 
Valerian, Sitka, 577 
Valeriana sitchensis, 560 
Vallisneria americana, 318 
Veery, 232 
Venerupis philippinarum, 78 
Venusia cambrica, 539 
pearsalli, 539 
Verbascum thapsus, 300 
Verbena bonariensis, 614 
Vermivora celata, 381 
chrysoptera, 232 
peregrina, 48 
Vernalgrass, Sweet, 177 
Verrucaria sp., 406 
amphibia, 411 
degelii, 411 
durietzii, 420 
epimaura, 406 
erichsenii, 410 
halizoa, 424 
maura, 408 
mucosa, 410 
sandstedei, 406 
schofieldii, 406 
silicicola, 424 
striatula, 406 
Vervain, Purpletop, 614 
Vetch, American, 300,574 
Common, 174 
Viburnum, 500 
cassinoides, 253 
Vicia americana, 300,560 
sativa, 174 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Villemure, M. and H. Jolicoeur. First Confirmed Occurrence 
of a Wolf, Canis lupus, South of the St. Lawrence 
River in Over 100 Years, 608 
Viola praemorsa ssp. praemorsa, 175 
renifolia var. brainerdii, 560 
Violet, Kidney-leaved, 574 
Yellow Montane, 175 
Vireo cassinii, 378 
flavifrons, 232 
gilvus, 232 
olivaceus, 231 
solitarius, 48 
Vireo, Blue-headed, 48 
Cassin’s, 378 
Red-eyed, 231 
Warbling, 232 
Yellow-throated, 232 
Vole, 342 
Heather, 438 
Long-tailed, 300 
Meadow, 342,400 
Montane, 300 
Prairie, 342 
Vole, Genus Phenacomys, in Alaska, The Heather, 438 
Vulpes vulpes, 40,58,69,96,244,249 273,368 


Wade, K.D., 95 
Wall-creeper, 614 
Warbler, Bay-breasted, 48 
Black-and-white, 230 
Black-throated Green, 48,232 
Blackburnian, 48 
Cerulean, 229 
Chestnut-sided, 232 
Golden-winged, 232 
Magnolia, 48 
Myrtle, 48 
Orange-crowned, 381 
Tennessee, 48 
Townsend’s, 382 
* Yellow, 86,232 
Yellow-rumped, 48,86,232,378 
Warbler, Dendroica cerulea Management on Eastern Ontario 
Bird Species, Predicting the Effects of Cerulean, 
229 
WARS 1983-2002 Special Annual Report Wildlife Accident 
Reporting and Mitigation, 645 
Wart-cress, Lesser, 613 
Washburn, B.E., 185 
Wasp, Parasitoid, 122 
Wasps, Baeus sp. and Gelis sp., in the Egg Sacs of the Wolf 
Spiders Pardosa moesta and Pardosa sternalis (Ara- 
neae, Lycosidae) in Southeastern Idaho, Occurrence 
of Parasitoid, 122 
Water-hemlock, Spotted, 574 
Watersnake, Northern, 435 
Watersnake, Nerodia sipedon sipedon, Frequency of Tail 
Breakage of the Northern, 435 
Waterthrush, Northern, 232 
Waterweed, Canada, 318 
Waxwing, Cedar, 232,378 
Weasel, 96,306 
Least, 403 
Weathervane X Japanese Scallop Hybrid, 78 
Weed, Willow, 568 


2004 


Weevil, White Pine, 41 
Wetzel, S., 195 
Whale, Dwarf Sperm, 495 
Pygmy Sperm, 495 
Sperm, 495 
Whale, Kogia breviceps, in the Northern Gulf of St. Law- 
rence, Canada, Stranding of a Pygmy Sperm, 495 
Wheatgrass, Crested, 300 
Whip-poor-will, 230 
Whitlow-grass, Wood, 572 
Wilkerson, C.D., 267 
Williston, P., Review by, 290 
Willow, 57,211,247,253,268,356,370,530 
Peachleaf, 309 
Willson, M.F. and S.M. Gende. Seed Dispersal by Brown 
Bears, Ursus arctos, in Southeastern Alaska, 499 
Willson, M.F. Gulls, Larus spp., Foraging at Pink Salmon, 
Oncorhynchus gorbuscha, Spawning Runs, 442 
Wilson, S.F., A. Hahn, A. Gladders, K.M.L. Goh, and 
D.M. Shackleton. Morphology and Population Char- 
acteristics of Vancouver Island Cougars, Puma con- 
color vancouverensis, 159 
Wintergreen, 196 
Wise, I.L., 309 
Wockia asperipunctella, 536 
Wolf, 115,127,134,278,368,603,608 
Eastern, 608 
Gray [Grey], 42,57,61,95,194,247,350,361,608 
Wolf, Canis lupus, Pack Density, Record High, 127 
Wolf, Canis lupus, South of the St. Lawrence River in Over 
100 Years, First Confirmed Occurrence of a, 608 
Wolverine, 56,61 
Wolverine, Gulo gulo luscus, Resting Sites and Caching 
Behavior in the Boreal Forest, 61 
Wolverines, Gulo gulo luscus, in the boreal forest, Effects 
of mid-winter snow depth on stand selection by, 56 
Wolves, Canis lupus, at Katmai National Park and Preserve, 
Alaska, Interactions of Brown Bears, Ursus arctos, 
and Gray, 247 
Wolves, Canis lupus, in and near Pukaskwa National Park, 
Ontario, Demographic patterns and limitations of Grey, 
95 
Wolves, Canis lupus, in Relation to Forest Harvesting in 
West-central Alberta, Winter Habitat Use by, 368 
Wolves, Canis lupus, Unusual Behavior by Bison, Bison 
bison, Toward Elk, Cervus elaphus, and, 115 
Wood-Peewee, Eastern, 232 
Woodlouse, 388 
Woodpecker, Downy, 358 
Woodsia alpina, 560 
ilvensis, 560 
Woodsia, Northern, 560 
Rusty, 560 
Woolly-heads, Slender, 169 
Tall, 169 


INDEX TO VOLUME 118 


675 


Woolly-heads, Psilocarphus elatior, an Endangered Herb in 
Canada, Conservation Evaluation of the Pacific Pop- 
ulation of Tall, 169 

Wren, Bewick’s, 378 

House, 378 

Wright, J.D. and J. Ernst. Effects of mid-winter snow depth 
on stand selection by Wolverines, Gulo gulo luscus, 
in the boreal forest, 56 

Wright, J.D. and J. Ernst. Wolverine, Gulo gulo luscus, Rest- 
ing Sites and Caching Behavior in the Boreal Forest, 
61 

Wright, K.G., 264 


Xanthia tatago, 542 
Xanthocephalus xanthocephalus, 207 
Xanthorhoe abrasaria, 539 
decoloraria, 539 
ferrugata, 539 
fossaria, 539 
iduata, 539 
lacustrata, 539 
Xanthoria, 420 
candelaria, 411 
parietina, 420 
Xanthotype sospeta, 538 
Xenolechia aethiops, 534 
Xestia normaniana, 543 
smithii, 543 
Xylena curvimacula, 542 
Xylotype arcadia, 542 


Yarrow, 166 

Yellowthroat, Common, 86,232 

Yew, Canada, 41 

Yponomeutidae, 532 

Ypsolopha canariella, 533 

dentiferella, 533 

Yukon Territory, New Records of Cyperaceae and Juncaceae 
from the, 266 

Yukon Territory VI, New Records of Vascular Plants in the, 
558 

Yunger, J.A., 341 


Zanclognatha lutalba, 540 
Zapus hudsonius, 403 
Zea mays, 253,365 
Zeiraphera canadensis, 536 
fortunana, 536 
unfortunana, 536 
Zenophleps alpinata, 539 
Zonotrichia leucophrys, 378 
Zophodia grossulariella, 536 
Zostera marina, 225 
Zostera marina, Wrack by Three Species of Ladybird Beetles 
(Coleoptera: Coccinellidae) in Prince Edward Island, 
Use of Eelgrass, 225 


676 


Index to Book Reviews 


Botany 

Berkutenko, A.N., H.G. Lumsden, and D. Lumsden. Flora 
and Climatic Conditions of the North Pacific: A Col- 
lection of Scientific Papers, 637 

Beresford-Kroeger, D. Arboretum America, A Philosophy of 
the Forest, 474 

Boulet, B. Les champignons des arbres de l’est de |’ Amerique 
du Nord, 146 

Clark, L.J. Lewis Clarks’s Field Guide to Wild Flowers of 
the Sea Coast in the Pacific Northwest, 476 

Clark, L.J. Wild Flowers of Field & Slope in the Pacific 
Northwest, 289,476 

Clark, L.J. Wild Flowers of Forest & Woodland in the Pacific 
Northwest, 289,476 

Clark, L.J. Wild Flowers of the Mountains in the Pacific 
Northwest, 289,476 

DiGregorio, M.J. and J. Wallner. Cape Cod Wildflowers: A 
Vanishing Heritage, 475 

Gleason, H.A. and A. Cronquist. Manual of Vascular Plants 
of Northeastern United States and Adjacent Canada: 
Second Edition, 147 

Martin, N.D. and N.M. Martin. Biotic Forest Communities 
of Ontario, 146 

Martin, P. The Wild Orchids of North America, North of Mex- 
ico, 147 

Nicholls G. and R. Lupp. Alpine Plants of North America: An 
Encyclopedia of Mountain Flowers from the Rockies 
to Alaska, 474 

Trelawny, J.G. Wild Flowers of the Yukon, Alaska & North- 
western Canada, 2nd Edition, 289 

Waldron, G. Trees of the Carolinian Forest: A Guide to 
Species, Their Ecology and Uses, 148 


Environment 

Boelens, B. and M. Watkins. Whose Bird? 150 

Bryson, M.A. Visions of the Land: Science, Literature, and the 
American Environment from the Era of Exploration 
to the Age of Ecology, 478 

Conner, J.K. and D.L. Hartl. A Primer of Ecological Genetics, 
480 

Czajkowski, C. Snowshoes & Spotted Dick; Letters from a 
Wilderness Dweller, 151 

Dietrich, W. Natural Grace: The Charm, Wonder, & Lessons 
of Pacific Northwest Animals & Plants, 152 

Frelich, L.E. Forest Dynamics and Disturbance Regimes: Stud- 
ies from Temperate Evergreen-Deciduous Forests, 150 

Josephson Weddell, B. Conserving Living Resources in the 
Context of a Changing World, 477 

Mitchell, B. Resource and Environmental Management in 
Canada: Addressing Conflict and Uncertainty, 640 

Newell, J. The Russian Far East, 638 

Suzuki, D. and Dressel, H. Good News for a Change: How 
Everyday People are Helping the Planet, 149 

Wolfe, D.W. Tales from the Underground: A Natural History 
of Subterranean Life, 153 

Young, A.G. and G.M. Clarke. Genetics, Demography and via- 
bility of Fragmented Populations, 478 


Miscellaneous 
Burnett, J.A. A Passion for Wildlife: The History of the Cana- 
dian Wildlife Service, 290 


THE CANADIAN FIELD-NATURALIST 


Vol. 118 


Edlow, J.A. Bull’s Eye: Unraveling the Medical Mystery of 
Lyme Disease, 641 

Jennings, J. The Canoe: A Living Tradition, 290 

Kareiva, P. and S.A. Levin. The Importance of Species: Per- 
spective on Expendability and Triage, 641 

Snetsinger, P. Birding on Borrowed Time, 291 

Yochelson, E.L. Smithsonian Institution Secretary Charles 
Doolittle Walcott, 481 


Zoology 

Attenborough, D. The Life of Mammals, 286 

Bird, D.M. The Bird Almanac, 630 

Bowman, T.D. Field Guide to Bird Nests and Eggs of Alaska’s 
Coastal Tundra, 632 

Bumstead, P.E. Canadian Skin and Scales: A Complete Ency- 
clopedia of Canadian Amphibians and Reptiles, 467 

Cannings, R. and S. Cannings. British Columbia: A Natural 
History. Revised and Updated, 631 

Carroll, D. Self-Portrait With Turtles: A Memoir, 471 

Cox, B. Conversations with an Eagle, 283 

Crump, M. In Search of the Golden Frog, 282 

del Hoyo, J., A. Elliott and D. Christie. Handbook of Birds of 
the World: Volume 9 Cotingas to Pipits and Wagtails, 
631 

Ducey, J.E. Birds of the Untamed West: The History of Bird- 
life in Nebraska, 1750 to 1975, 466 

Duff, A. and A. Lawson. Mammals of the World: A Checklist, 
634 

Dunne, P. Pete Dune on Bird Watching: The How-to, Where- 
to and When-to of Birding, 473 

Early, C. Sparrows and Finches of the Great Lakes and Eastern 
North America, 288 

Early, C. Warblers of the Great Lakes and Eastern North Amer- 
ica, 288 

Eisner, T. For Love of Insects, 468 

Feldhamer, G.A., B.C. Thompson and J.A. Chapman. Wild 
Mammals of North America: Biology, Management, 
and Conservation (Second edition), 472 

Ferrari, A. & A. Sharks, 282 

Ferri, V. Turtles and Tortoises, 143 

Heide-Jérgensen, M.P., O. Wiig, and D.G. Pike. Belugas in 
the North Atlantic and the Russian Arctic, 144 

Heintzelman, D. All-Weather Hawk Watcher’s Field Journal, 
471 

Heintzelman, D.S. Guide to Hawk Watching in North Amer- 
ica, 632 

Johnsgard, P.A. North American Owls, Biology and Natural 
History, Second Edition, 284 

Johnston, D.W. The History of Ornithology in Virginia, 470 

Jones, H.L. Birds of Belize, 140 

Joynt, A. and M.G. Sullivan. Fish of Alberta, 281 

Limbert, M. The Uses and Curation of Birds’ Egg Collec- 
tions: An Examination and Biography, 465 

Linder, G., S.K. Krest, D.W. Sparling. Amphibian Decline: 
An Integrated Analysis of Multiple Stressor Effects, 
138 

Lockwood, J.A. Locust: The Devastating Rise and Mysterious 
Disappearance of the Insect that Shaped the American 
Frontier, 633 

McCabe, R.E., B.W. O’ Gara, and H.M. Reeves. Prairie Ghost: 
Pronghorn and Human Interaction in Early America, 
635 


2004 


McCulloch, N. A Guide to the Birds of St. Helena and Ascen- 
sion Island, 636 

Melvin, E.F. and J.K. Parrish. Seabird Bycatch: Trends, Road- 
blocks, and Solutions, 285 

Morton, M.L. The Mountain White-Crowned Sparrow: Migra- 
tion and Reproduction at High Altitude, 284 

O'Toole, C. The Firefly Encyclopedia of Insects and Spiders, 
145 

Perrins, C. The Firefly Encyclopedia of Birds, 139 

Pianka, E.R., L.J. Vitt. Lizards: Windows to the Evolution 
of Diversity, 141 

Pough, F.H., R.M. Andrews, J.E. Cadle, M.L. Crump, A.H. 
Savitsky and D.D. Wells. Herpetology: Third Edition, 
140 

Richards, J.M., Y.T. Tymstra and A.W. White. Birds of Nuna- 
vut: A Checklist, 465 

Rising, J.D. Geographic Variation in Size and Shape of Svan- 
nah Sparrows (Passerculus sanwichensis), 287 

Sharpe, R.S., W.R. Silcock and J.G. Jorgensen. Birds of Neb- 
raska: Their Distribution and Temporal Occurrence, 
466 


INDEX TO VOLUME 118 


677 


Simpson, K. and N. Day. Birds of Australia (Seventh edition), 
630 

Sinclair, P.H., W.A. Nixon, C.D. Eckert and N.L. Hughes. 
Birds of the Yukon Territory, 288 

Somma, L.A. Parental Behavior in Lepidosaurian and Tes- 
tudinian Reptiles: A Literature Survey, 635 

Stewart, K.W. and D.A. Watkinson. The Freshwater Fishes 
of Manitoba, 469 

Stuart, C. & T. Birds of Africa: From Seabirds to Seed-eaters, 
138 

Swarth, C.W., W.M. Roosebrug and E. Kiviat. Conservation 
and Ecology of Turtles of the Mid-Atlantic Region: 
A Symposium, 468 

Waldbauer, G. What Good are Bugs? Insects in the Web of 
Life, 144 

Wheeler, B. Raptors of Eastern North America, 143 

Wheeler, B.K. Raptors of Western North America, 142 


Advice for Contributors to The Canadian Field-Naturalist 


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678 


| TABLE OF CONTENTS (concluded) Volume 118 Number 4 


| Tributes 
| A tribute to Thomas Henry Manning 1911-1998 BRENDA CARTER 


| A tribute to Victor Kent Prest 1913-2003 A. S. Dyke, D. A. Hopcson, E. L. BOUSFIELD, R. E. BEDFORD 


‘Book Reviews 

| ZOOLOGY: Birds of Australia (Seventh edition) — The Bird Almanac — Handbook of Birds of the World: 
Volume 9 Cotingas to Pipits and Wagtails — British Columbia: A Natural History Revised and Updated 
—A Field Guide to Bird Nests and Eggs of Alaska’s Coastal Tundra — Guide to Hawk Watching in 
North America — Locust: the Devastating Rise and Mysterious Disappearance of the Insect that shaped 
the American Frontier — Mammals of the World: A Checklist — Parental Behavior in Lepidosaurian 
and Testudinian Reptiles: A Literature Survey — Prairie Ghost: Pronghorn and Human Interaction in 
Early America — A Guide to the Birds of St. Helena and Ascension Island 


| BoTaNy: Flora and Climatic Conditions of the North Pacific: A Collection of Scientific Papers 


| ENVIRONMENT: The Russian Far East — Resource and Evironmental Management in Canada: 
Addressing Conflict and Uncertainty 


| MISCELLANEOUS: Bull’s Eye: Unraveling the Medical Mystery of Lyme Disease; The Importance of Species: 
| Perspective on Expendibility and Triage 


q 
| NEw TITLES 


News and Comment 
| RENEW 2004 Recovery of Nationally Endangered Wildlife in Canada Annual Report (14) — Calendar: 
| Endangered Reptiles of Canada 2004 and 2005 — Canadian Species at Risk November 2004 — 
Froglog: Newsletter of the Declining Amphibian Populations Task Force (65, 66) — The Boreal Dip 
Net/L’Epuisette Boreale: Newsletter of the Canadian Amphibian and Reptile Conservation Network/ 
Reseau Canadien de Conservation des Amphibiens et des Reptiles 9(1) December 2004 — Marine Turtle 
Newsletter (106) — WARS 1983-2002 Special Annual Report Wildlife Accident Reporting and Mitigation 


The Ottawa Field-Naturalists’ Club Awards for 2003 
Errata The Canadian Field-Naturalist 118(3) 
| Index to Volume 118 COMPILED BY LESLIE CODY 


| Advice to Contributors 


| Mailing date of the previous issue 118(3): 21 October 2005 


2004 


618 
626 


630 
637 


638 


641 
643 


644 


646 
649 
650 
678 


THE CANADIAN FIELD-NATURALIST Volume 118 Number 4 


2004) 


Articles 


Stranding of a Pygmy Sperm Whale, Kogia breviceps, in the northern Gulf of St. Lawrence, Canada 


LENA MEASURES, BENOIT ROBERGE, and RICHARD SEARS 


Seed dispersal by Brown Bears, Ursus arctos, in southeastern Alaska 
Mary F. WILLSON and Scott M. GENDE 


A mapping of the present and past forest-types of Prince Edward Island 
D. G. SoBey and W. M. GLEN 


Inventaire printanier d’une frayere multispécifique: ’ichtyofaune des rapides de la riviere 
Gatineau, Québec ANNIE COMTOIS, FRANCOIS CHAPLEAU, CLAUDE B. RENAUD, 
HENRI FOURNIER, BRENT CAMPBELL, and RICHARD PARISEAU 


Moths and butterflies (Lepidoptera) of the boreal mixedwood forest near Lac La Biche, 
Alberta, including new provincial records 
GREG R. POHL, DAvID W. LANGOR, JEAN-FRANCOIS LANDRY, and JOHN R. SPENCE 


Pilose Braya, Braya pilosa Hooker (Cruciferae, Brassicaceae), an enigmatic endemic of 
arctic Canada JAMES G. HARRIS 


New records of vascular plants in the Yukon Territory VI 
WILLIAM J. Copy, CATHERINE E. KENNEDY, BRUCE BENNETT, and PHIL CASWELL 


Hybridization between a Green Turtle, Chelonia mydas, and a Loggerhead Turtle, Caretta caretta, 


and the first record of a Green Turtle in Atlantic Canada 
MICHAEL C. JAMES, KATHLEEN MARTIN, and PETER H. DUTTON 


A snow-tracking protocol used to delineate local Lynx, Lynx canadensis, distributions 
JOHN R. SQUIRES, KEVIN S. MCKELVEY, and LEONARD F. RUGGIERO 


Hummock vegetation at the arctic tree-line near Churchill, Manitoba JORG TEWS 


Effects of Lodgepole Pine Dwarf Mistletoe, Arceuthobium americanum, on Jack Pine, 
Pinus banksiana, growth in Manitoba Brock Epp and JACQUES C. TARDIF 


Notes 
Excavation of an Arctic Fox, Alopex lagopus, den by a Polar Bear, Ursus maritimus 
EVAN RICHARDSON and RYAN K. BROOK 


Two Great Black-backed Gulls, Larus marinus, kill male Longtailed Duck, Clangula hyemalis 
MARTHA DowsLey and ANDREW CIRTWILL 


First record of Mountain Lions, Puma concolor, in Elk Island National Park, Alberta 
GLYNNIS A. Hoop and TIM NEUFELD 


First confirmed occurrence of a Wolf, Canis lupus, south of the St. Lawrence River in over 100 years 
Mario VILLEMURE and HELENE JOLICOEUR 


Is cost of locomotion the reason for prolonged nesting forays of Snapping Turtles, Chelydra serpentina? 
SHANE R. DE SOLLA and KIM J. FERNIE 


Significant vascular plant records from the Hamilton area, Ontario CARL ROTHFELS 


Do Rhinoceros Auklet, Cerorhinca monocerata, fledglings fly to the sea from their natal burrows 
JAMES L. HAYWARD and JERE K. CLAYBURN 


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sas pei . Pericto 


Diamar rates: 
wed 


sabe tae +3 
whens 
gh res 


AAiNes 


Sa sgh dare Wears 


Ae ita eaae ; ; 
preg sberse swear M om hd x zg raha 2a aL 


ire ele u $ ‘i Apron con i at c Phir eran 
a iw Lives ia 2 Fa MARY: ms 
(Cet pantie ae t : C ‘ LY ea ck ype we 2 Lol ie